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Akademiya Nauk SSSR 



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FLORA of the U.S. S.R. 



Volume IX 



V. L. Komarov, Editor 



Rosales and Sarraceniales 



TRANSLATED FROM RUSSIAN 



Published for the Smithsonian Institution 

and the National Science Foundation, Washington, D.C. 

by the Israel Piogram for Scientific Translations 



ill 

BOTANICHESKII INSTITUT AKADEMII NAUK SSSR 



Botanical Institute of the Academy of Sciences of the U.S.S.R. 

» 

FLORA OF THE U.S.S.R. 

(Flora SSSR) 



Volume IX 

Rosales and Sarraceniales 



Chief Editor Academician V.L. Komarov 
Volume Editor S.V. Yuzepchuk 



Compiled by 

A.G. Borisova, V.L. Komarov, A.N. Krishtofovich, 

A.S. Lozina-Lozinskaya, V.P. Maleev, I.V. Palibin, 

A.I. Poyarkova, Yu.D. Tsinzerling, 

and S.V. Yuzepchuk 




rS 



Izdatel'stvo Akademii Nauk SS£R 

Moskva -Leningrad ^^^/yOSf 



1939 



Translated from Russian 



Israel Program for Scientific Translations 
Jerusalem 1971 



TT 70-50110 

Published Pursuant to an Agreement with 

THE SMITHSONIAN INSTITUTION, U. S. A. 

and 

THE NATIONAL SCIENCE FOUNDATION, WASHINGTON, D. C. 



Copyright © 1971 

Israel Program for Scientific Translations Ltd. 

IPST Cat. No. 5773 



Translated by L. Behrman 
Edited by Z. Blake 



Printed in Jerusalem by Keter Press 
Binding: Wiener Bindery Ltd., Jerusalem 



Available from the 
U. S. DEPARTMENT OF COMMERCE 
National Technical Information Service 
Springfield, Va. 22151 



XII/7/3 



TABLE OF CONTENTS 



Systematic Index of Species in Volume IX 

Preface 

Order 22. Sarraceniales Engl 

Family LXXII. Droseraceae DC 

Genus 693. Aldrovanda Monti 

Genus 694. Drosera L 

Order 23. Rosales Lindl 

Family LXXIII. Crassulaceae DC 

Key to Genera 

Subfamily 1. Crassuloideae Berger 

Genus 695. Ti 11 a ea (Mich.) L 

Subfamily 2. Cotyledonoideae Berger 

Genus 696. Umbilicus DC 

Subgenus Chiastophyllum 

Subfamily 3. Sempervivoideae Berger 

Genus 697. Sempervivum L , 

Subfamily 4. Sedoideae Berger 

Genus 698. RhodiolaL. 

Genus 699. Sedum L , 

Genus 700. Pseudosedum (Boiss.) Berger 

Genus 701. Orostachys (DC.) Fisch. 

Genus 702. Ro sul a r ia (DC.) Stapf 

Subfamily 5. Penthoroideae (Engl.) A.Bor. 

Genus 703. Penthorum Gronov 

Family LXXIV. Saxifragaceae DC 

Subfamily 1. Saxifragoideae A.Br 

Tribe 1. Saxifrageae DC 135 

Genus 704. A s t i 1 b e Hamilton 

Genus 705. Bergenia Monch 

Genus 706. Saxifraga L. 

* [Russian page numbers appear in the left-hand margin of the text.] 



Russian 


English 


page* 


page 


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vii 


vi 


1 


1 


3 


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1 


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5 


6 


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9 


10 


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13 


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15 


14 


24 


20 


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45 


36 


99 


79 


108 


86 


114 


90 


132 


104 


132 


104 


134 


106 


135 


107 


135 


107 


135 


107 


136 


108 


138 


109 



Genus 707. Mi tell a L 

Genus 708. Chrysosplenium L 

Tribe 2. Parnassieae S.F. Gray 

Genus 709. Parnassia L 

Subfamily 2. Hydrangeoideae A. Br 

Tribe 1. Philadelpheae Rchb 

Genus 710. PhiladelphusL 

Genus 711. Deutzia Thbg 

Tribe 2. Hydrangeae DC 

Genus 712. Hydrangea L 

Subfamily 3. Ribesioideae Engl 

Genus 713. Ribes L 

Subgenus 1. Ribesia (Berl.) Jancz. 

Subgenus 2. Her i tier a Jancz 

Subgenus 3. Grossularioides Jancz. 
Subgenus 4. Euc ore osma Jancz. ... 

Subgenus 5. BerisiaSpach 

Genus 714. Grossularia Mill 

Family* Pittosporaceae Lindl 

Genus* Pi tt os po rum Dryand 

Family LXXV. Hamamelidaceae Lindl 

Genus 715. Parrotia C.A.M 

Family* Eucommiaceae Van-Tieghem 

Genus* Eucommia Oliv 

Family LXXVI. Platanaceae Lindl 

Genus 716. Platanus L 

Family LXXVII. Rosaceae Juss , 

Key to Subfamilies 

Subfamily 1. Spiraeoideae Agardh 

Genus 717. Phy so c a r pus Maxim. .... 

Genus 718. Spiraea L 

Subgenus 1. Protospiraea Nakai. 
Subgenus 2. Metaspiraea Nakai. 

Genus 719. Sibirae a Maxim , 

Genus 720. A r u n c u s Adans , 

Genus 721. Sorbaria A.Br 

Genus 722. Spiraeanthus Maxim. 

Genus 723. Exochorda Lindl 

Subfamily 2. Pomoideae Focke 

Genus 724. Cotoneaster Medik. ... 

Genus 725. CydoniaMill 

Genus 726. PyrusL 

Genus 727. MalusMill 

Genus 728. Sorbus L 



199 


154 


200 


155 


215 


166 


216 


166 


220 


169 


220 


170 


220 


170 


223 


172 


225 


174 


225 


174 


226 


175 


226 


175 


232 


179 


246 


189 


247 


190 


248 


191 


257 


198 


267 


205 


270 


208 


270 


208 


271 


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272 


209 


273 


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210 


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274 


211 


279 


215 


280 


216 


281 


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286 


220 


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224 


306 


235 


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243 


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244 


318 


245 


319 


246 


334 


257 


336 


259 


357 


275 


372 


286 



Subgenus 1. Eu-Sorbus Kom 

Subgenus 2. H a h n i a Medik 

Genus 729. Micromeles Decne 

Genus *Eriobotrya Lindl 

Genus 730. Amelanchier Medik 

Genus 731. Pyracantha Roem 

Genus 732. M e s p i 1 u s L 

Genus 733. Crataegus L 

Addenda VIII (Diagnoses of New Species Mentioned in Volume IX) 

Index Alphabeticus 

Vegetation Regions of the USSR 

List of Abbreviations 



372 


286 


387 


296 


406 


310 


407 


311 


408 


312 


413 


315 


414 


316 


416 


317 


469 


357 


511 


397 


541 


416 




419 



SUBJECTS AND CONTRIBUTORS 



Preface and Indexes 

Family Crassulaceae 

Subfamily Saxifragoideae 

Subfamilies Hydrangeoideae, Ribesioideae, 
Spiraeodieae. Genera Cotoneaster, Cydonia, 
Micro meles, Eriobotrya, Amelanchier, 
Pyracantha, Mespilus, Crataegus 

Families Pittosporaceae, Hamamelidaceae, 
Eucommiaceae, Platanaceae 

Genus P y r u s 

Genus M a 1 u s 

Subgenus Eu-Sorbus of Genus S or bus 

Subgenus Hah ni a of Genus Sorb us 

Reports on Discovery of Plant Fossils 

Addenda — Descriptiones plantarum novarum 
in tomo IX Florae URSS commemoratarum. 



Editorial Staff 

Arranged by A.G.Borisova 

Arranged by A.S.Lozina-Lozinskay£ 



Arranged by A.I.Poyarkova 

Arranged by I.V.Palibin 
Arranged by V.P.Maleev 
Arranged by S.V.Yuzepchuk 
Arranged by V.L.Komarov 
Arranged by Yu.D.Tsinzerling 
A.N.Krishtofovich 



The plates were drawn by the following artists: Z.V.Kobyletskaya — I— XI, XXII; 
A.S.Lozina-Lozinskaya — XII; S.A.Moiseeva - XIII, XV, XXVII; E.A.Derevitskaya - 
XIV; M.M.Parfenenko (deceased) - XVI; E.V.Blagoveshchenskaya - XVII-XIX, 
XXIII-XXVI, XXVIII-XXX; O.P.Voronova - XX, XXI. 



SYSTEMATIC INDEX OF SPECIES IN VOLUME IX* 



Order 22. Sarraceniales Rgl. 
Family LXXII. Droseraceae DC. 

Genus 693. Aldrovanda Monti 

Russian page** 

1. A. vesiculosa L j 

Genus 694. Drosera L. 

1. D. rotundifolia L. . . . 2 

2. D. anglica Huds 5 

3. D. intermedia Hayne g 

4. D. obovata Mert. et Koch 6 

Order 23. Ro sales Lindl. 

Family LXXIII. Crassulaceae DC. 

Subf am ily 1. Crassuloideae Berber 
Genus 695. Tillaea (Mich.) L. 

1. T. Vaillantii Willd 11 

2. T. aquatica L 12 

3- T. alata Viv 12 

Subfamily 2. Cotyledonoideae Berger 

Genus 696. Umbilicus DC 
1. U. oppositifolius Ldb 14 

Subfamily 3. Sempervivoideae Berger 
Genus 697. Sempervivum L. 
Section 1. Eu-Sempervivum Schoenl. 

1. S. pumilum M. B 16 

2. S. tectorum L 17 

3. S. caucasicum Rupr 18 

4. S. ruthenicum (C. Koch) Schnittsp. et Lehm 21 

5. S. globiferum L 21 

6. S. glabrifolium A. Bor 22 

Section 2. jovisbarba DC. 

7. S. soboliferum Sims 23 

* [This index has been reproduced photographically from the Russian original. J 
*• [Russian page numbers appear in the left-hand margin of the text.] 



Subfamily 4. Sedoideae Berger 
Genus 698. Rhodiola L. 
Section 1. Clementsia (Rose) A. Bor. 

1. R. Semenovii (Rgl. et Herd.) A. Bor 28 

Section 2. Eu-Rhodiola Schrenk 

2. R. rosea L 29 

3. R. arctica A. Bor 30 

4. R. iremelica A. Bor 31 

5. R. sachalinensis A. Bor 

6. R. heterodonta (Hook, et Thorns.) A. Bor 32 

— R. viridula A. Bor 500 

7. R. borealis A. Bor 32 

8. R. atropurpurea (Turcz.) Trautv. et Mey 

9. R. Kirilowii Rgl. et Maxim 34 

10. R. Hnearifolia (Rgl. et Schmalh.) A. Bor 35 

11. R. pinnatifida A. Bor 36 

12. R. Stephani (Cham.) Trautv. et Mey 36 

13. R. algida (Ldb.) Fisch. et Mey 37 

14. R. Komarovii A. Bor & 

Section 3. Chamae-Rhodiola Schrenk 

15. R. quadrifida (Pall.) Fisch. et Mey 39 

16. R. kaschgarica A. Bor 

17. R. pamiroalaica A. Bor 

18. R. coccinea (Royle) A. Bor 41 

1 9. R. gelida Schrenk 41 

20. R. reticulata A. Bor 

21. R. Litvinovii A. Bor 45 

Genus 699. Sedum L. 
Section 1. Telephium S. T. Gray 

CO 

1. S. purpureum (L.) Schult 

2. S. mugodscharicum A. Bor 

3. S. parvistamineum V. Petrov 

4. S. maximum (L.) Suter 

5. S. telephium L * 

6. S. caucasicum (Grossh.) A. Bor 

7. S. verticillatum L 

8. S. viviparum Maxim 

9. S. ussuriense Kom 

10. S. pallescens Freyn 

11. S. eupatorioides Kom 

12. S. alboroseum Baker 

13. S. Ewersii Ldb 

14. S. pluricaule Kudo 

15. S. cyaneum Rud 

e i 66 

— S. anacampseros L 

— S. fabaria Koch 

Section 2. Populisedum Berger 

16. S. populifolium Pall 



Section 3. Aizoon Koch 

17. S. aizoon L. 57 

18. S. hyperaizoon Kom 63 

19. S. kamtschaticum Fisch 69 

20. S. Middendorfianum Maxim 69 

21. S. hybridum L 70 

22. S. litorale Kom 71 

23. S. Selskianum Rgl. et Maak 71 

Section 4. S e da Genuina Koch 

24. S. Stevenianum Rouy et Camus 72 

25. S. stoloniferum S. G. Gmel 73 

26. S. spurium M. B 74 

27. S. oppositifolium Sims 77 

28. S. involucratum M. B 78 

29. S. obtusifolium C. A. M 79 

30. S Listoniae Vis 79 

31. S. tenellum M. B 80 

32. S. lydium Boiss 81 

33. S. album L 82 

34. S. gracile C. A. M 83 

35. S. lenkoranicum Grossh 84 

36. S. subulatum (C.A. M.) Boiss 84 

37. S. Alberti Rgl 85 

38. S. reflexum L 86 

39. S. sexangulare L 87 

40. S. acre L 88 

41. S. polytrichoides Hemsl 89 

Section 5. Epeteium Boiss. 

42. S. hispanicum L 89 

43. S. pentapetalum A. Bor 90 

44. S. bucharicum A. Bor. 91 

45. S. pallidum M. B 92 

46. S. corymbosum Grossh 93 

47. S. atratum L 93 

48. S. villosum L 94 

49. S. annuum L 95 

50. S. nanum Boiss 95 

51. S. rubens L 96 

52. S. rubrum (L.) Thell 97 

53. S. tetramerum Traulv 98 

54. S. aetnense Tin 98 

Genus 700. Pseudosedum (Boiss.) Berger 
Section 1. Li e v e ni a A. Bor. 

1. P. Lievenii (Ldb.) Berger 101 

2. P. longidentatum A. Bor 102 

3. P. condensatum A. Bor 102 

4. P. bucharicum A. Bor 105 

Section 2. Tuberaria A. Bor. 

5. P. ferganense A. Bor 105 



Section 3. Campanella A. Bor. 

6. P. multicaule (Boiss et Buhse) A. Bor 106 

7. P. Fedtschenkoanum A. Bor 107 

8. P. campanuliflorum A. Bor 107 

9. P. karatavicum A. Bor 108 

Genus 701. Orostachys (DC.) Fisch. 
Section 1. Euappendiculata A. Bor. 

1. O. malacophylla (Pall.) Fisch 110 

Section 2. Appendiculata A. Bor. 

2. O. spinosa (L.) C. A. M. . . . 110 

3. O. thyrsiflora Fisch ■ 112 

4. O. cartilaginea A. Bor 112 

5. O. fimbriata (Turcz.) Berger 113 

Genus 702. Rosularia (DC.) Stapf 
Section 1. Sempervivoides Boiss 

1. R. pilosa (M. B.) A. Bor 117 

2. R. sempervivoides (Fisch.) A. Bor 118 

Section 2. Eu-Rosularia Berger 

3. R. sempervivum (M. B.) Berger 119 

4. R. radiciflora (Steud.) A. Bor 120 

5. R. persica (Boiss.) Berger 120 

6. R. elymaitica (Boiss. et Hausskn.) Berger 121 

7. R. Lipskyi A. Bor 121 

8. R. hissarica A. Bor • 122 

Section 3. Orientalia A. Bor. 

9. R. paniculata (Rgl. et Schmalh.) Berger 123 

10. R. subspicata (Freyn et Sint.) A. Bor. 123 

11. R. glabra (Rgl. et Wink! ) Berger 124 

12. R turkestanica (Rgl. et Winkl.) Berger 127 

Section 4. Campanella A. Bor. 

13. R. kokanica (Rgl. et Schmalh.) A. Bor 128 

14. R. lutea A. Bor 128 

15. R. alpestris (Kar. et Kir.) A. Bor 129 

16. R. Schischkinii A. Bor 130 

17. R. tadzhikistana A. Bor • 130 

18. R. platyphylla (Schrenk) Berger 131 

Subfamily 5. Penthoroideae Engl. 
Genus 703. Penthorum Gronow 

1. P. chinense Pursh • 132 

2. P. humile Rgl. et Maack 133 

Family LXXIV. Saxifragaceae DC. 

Subfamily 1. Saxifragoidcae A. Br. 
Tribe 1. SAXIFRAGEAE DC. 
Genus 704. Astilbe Hamilton 

1. A. chinensis (Maxim.) Franch. et Sav 135 



Genus 705. Bergenia MoncVi 

1. B. crassifolia (L.) Fritsch 137 

2. B. pacifica Kom 137 

Genus 706. Saxifraga L. 
Section 1. Boraphylla Engl. 

1. S. nudicaulis D. Don 145 

2. S. Sieversiana Sternb 146 

3. S. Korshinskyi Kom 146 

4. S. reniformis Ohwi 147 

5. S. punctata L 147 

6. S. Nelsoniana D. Don 148 

7. S. purpurascens Kom 149 

8. S. manshuriensis (Engl.) Kom 149 

9. S. astilbeoides A. Los 149 

10. S. Redowskiana Sternb 150 

11. S. dahurica Willd 151 

12. S. grandipetala (Engl, et Irmsch.) A. Los 151 

13. S. calycina Sternb 151 

14. S. nivalis L 152 

15. S. tenuis (Wahlb.) Sm 155 

16. S. Tilingiana Rgl. et Til • ■. 155 

17. S. melaleuca Fisch. • 156 

18. S. sachalinensis Fr. Schmidt 156 

19. S. hieracifolia W. et K 157 

20. S. foliolosa R. Br 157 

21. S. Redowskii Adams 158 

22. S. stellaris L 158 

23. S. Merkii Fisch 159 

Section 2. Hirculus (Haw.) Tausch 

24. S. parnassioides Rgl. et Schmalh 160 

25. S. hirculus L 160 

26. S. flagellars Willd • 161 

27. S. Komarovii A. Los 162 

28. S. setigera Pursh 162 

29. S. serpyllifolia Pursh 163 

Section 3. Miscopetalum Haw. 

30. S. coriifolia (Somm. et Lev.) Grossh 163 

Section 4. Cymbalaria Griseb. 

31. S. Huetiana Boiss 164 

32. S. cymbalaria L 165 

Section 5. Tridactylites Haw. 

33. S. tridactylites L . 165 

34. S. adscendens L 166 

Section 6. Nephrophyllum Gaud. 

35. S. irrigua M. B 167 

36. S. granulata L 167 

37. S. cernua L 168 



38. S. sibirica L 171 

39. S. mollis Smith 172 

40. S. rivularis L 172 

41. S. bracteata D. Don 173 

42. S. exilis Stephan 173 

Section 7. Dactyloides Tausch. 

43. S. androsacea L 174 

44. S. sileniflora Sternb 174 

45. S. lactea Turcz 175 

46. S. caespitosa L 175 

47. S. terektensis Bge 176 

48. S. moschata Vulf 177 

49. S. pontica Albow 178 

50. S. exarata Vill 178 

51. S. adenophora C. Koch 179 

52. S. verticillata A. Los 180 

Section 8. Trachyphyllum Gaud. 

53. S. anadyrensis A. Los 180 

54. S. bronchialis L 183 

55. S. spinulosa Adans 183 

56. S. firma Litw 184 

57. S. cherlerioides Don 185 

58. S. Eschscholtzii Sternb 185 

Section 9. Xanthizoon Griseb. 

59. S. aizoides L • 186 

Section 10. Euaizoonia (Schott) Engl. 

60. S. cartilaginea Willd 186 

61. S. Kolenatiana Rgl 187 

Section 11. Kabschia Engl. 

62. S. juniperifolia Adams 188 

63. S. subverticillata Boiss 188 

64. S. colchica Albow 189 

65. S. laevis M. B 189 

66. S. pseudolaevis Oerting. . 190 

67. S. scleropoda Somm. et Lev 190 

68. S. abchasica Oetting 191 

69. S. caucasica Somm. et Lev 191 

70. S. Desoulavyi Oetting 192 

71. S. Kusnezowiana Oetting " ■ 192 

72. S. carinata Oerting 193 

73. S. Kotschyi Boiss 193 

74. S. Albertii Rgl. et Schmalh 194 

75. S. columnaris Schmalh 194 

76. S. Dinnikii Schmalh 197 

Section 12. Porphyrion Tausch. 

77. S. oppositifolia L 197 



Section 13. D i p t er a(Borkh.) Engl, et Irmsch. 

78. S. cortusifolia Sieb. et Zucc 198 

79. S. oblongifolia Nakai 198 

Genus 707. Mitella L. 

1. M. nuda L 199 

Genus 708. Chrgsosplenium L. 

1. Ch. nudicaule Bge 202 

2. Ch. peltatum Turcz 202 

3. Ch. alternifolium L • 203 

4. Ch. tetrandrum (Lund) Th. Fries • • . . 204 

5. Ch. beringianum Rose 204 

— Ch. Wrightii Franch. et Sav 204 

6. Ch. flagelliferum Fr. Schmidt 205 

7. Ch. filipes Kom 206 

8. Ch. Komarovii A. Los 206 

9. Ch. Sedakowii Turcz 207 

10. Ch. ovalifolium M. B 207 

11. Ch. thianschanicum Krassn 208 

12. Ch. ramosum Maxim 208 

13. Ch. pilosum Maxim 209 

14. Ch. sinicum Maxim 210 

15. Ch. kamtschaticum Fisch 210 

16. Ch. trachyspermum Maxim 213 

17. Ch. rimosum Kom 214 

18. Ch. dubium J. Gay 214 

19. Ch. baicalense Maxim 215 

Tribe 2. PARNASSIEAE S. F. GRAY 
Genus 709. Parnassia L. 

1. P. palustris L 216 

2. P. Kotzebuei Cham, et Schlecht. 218 

3. P. bifolia Nekras 218 

4. P. Laxmanni Pall 219 

Subfamily 2. HydrangeoideaeR. Br. 
Tribe 1. PHILADELPHEAE RCHB. 
Genus 710. Philadelphus L. 

1. Ph. caucasicus Koehne 220 

2. Ph. tenuifolius Rupr. et Maxim 221 

3. Ph. Schrenkii Rupr. et Maxim 222 

— Ph. latifolius Schrad 223 

Genus 711. Deutzia Thbg. 

1. D. amurensis (Rgl.) Airy-Show 224 

2. D. glabrata Kom . 224 

Tribe 2. HYDRANGEAE DC. 

Genus 712. Hydrangea L. 

1. H. petiolaris Sieb. et Zucc 225 

2. H. paniculata Sieb 226 



Subfamily 3. Ribesioideae Engl. 

Genus 713. Ribes L. 
Subgenus 1. Ribesia (Berl.) Jancz. 

1. R. triste Pall • . 232 

— R. vulgare Lam 233 

2. R. manschuricum Kom 234 

3. R. Palczewskii A. Pojark 237 

4. R. rubrum L • 237 

5. R. hispidulum A. Pojark 238 

6. R. pubescens Hedl 239 

— R. scandicum Hedl 240 

7. R. Meyeri Maxim 241 

8. R. latifolium Jancz 242 

9. R. pallidiflorum A. Pojark 242 

10. R. altissimum Turcz 243 

11. R. atropurpureum C. A. M. . 244 

12. R. Biebersteinii Berl 245 

Subgenus 2. Heritiera Jancz. 

13. R. sachalinense Nakai 246 

14. R. malvifolium A. Pojark. -. • 246 

Subgenus 3. Grossularioides Jancz. 

15. R. horridum Rupr 247 

Subgenus 4. Eucoreosma Jancz. 

16. R. ussuriense Jancz. . ; 248 

17. R. pauciflorum Turcz 248 

18. R. kolymense Kom 251 

19. R. turbinatum A. Pojark 252 

20. R. Janczewskii A. Pojark 252 

21. R. nigrum L 252 

22. R. dikuscha Fisch 254 

23. R. procumbens Pall 255 

24. R. fragrans Pall 256 

25. R. graveolens Bge 256 

Subgenus 5. B e r i s i a Spach. 

26. R. orientale Desf 257 

27. R. melananthum Boiss. et Hoh 258 

28. R. heterotrichum C. A. M • • • • 258 

29. R. villosum Wall ■ 259 

30. R. Maximowiczianum Kom 260 

31. R. aipinum L 260 

32. R. Komarovii A. Pojark 263 

33. R. lucidum Kit 264 

34. R. diacantha Pall 264 

35. G. saxatile Pall 265 

36. R. pulchellum Turcz 266 

— R. sanguineum Pursh 266 

— R. aureum Pursh 266 



Genus 714. Grossularia Mill. 

1. G. reclinata (L.) Mill 26g 

2. G. acicularis (Smith) Spach 2fiQ 

3. G. burejensis (Fr. Schmidt) Berger 970 

Family * Pittosporaceae Lindl. 
Genus * PUtosporam Dryand. 

— P. tobira Dryand. . « 7 -j 

Family LXXV. Hamamelidaceae Lindl. 
Genus 715 . Parrotia C. A. M. 
1. P. persica (D. C.) C.A, M 272 

Family * Eucommiaceae Van-Tieghem 
Genus * Eucommia Oliv. 

— E. ulmoides Oliv 273 

Family LXXVI. Platanaceae Lindl. 
Genus 716. Platanus L. 
Section 1- Orientalis Dode 

— P. orientalis L 275 

— P. cuneata W 275 

1. P. digitata Gordon 276 

2. P. orientalior Dode • 276 

Section 2. Occidentalis Dode 

— P. occidentalis L 279 

Family LXXVII. Rosaceae Juss. 

Subfamily 1. Spiraeoideae Agardh 

Genus 717. Physocarpus Maxim. 

1. Pb. amurensis Maxim 282 

2. Ph. ribesifolia Kom 283 

— Ph. opulifolia (L.) Maxim 283 

Genus 718. Spiraea L. 
Subgenus 1. Protospiraea Nakai 
Section 1. Spiraria Ser. 

1. S. salicifolia L 286 

2. S. humilis A. Pojark 287 

Section 2. Calospira C. Koch 

3. S. betulifolia Pall 288 

4. S. Beauverdiana C. K. Schn 289 

5. S. baldshuanica B. Fedtsch 290 

Subgenus 2. Metaspiraea Nakai 
Section 1- Chamaedryon Ser. 

6. S. chamaedryfolia L 291 

7. S. flexuosa Fisch 291 



8. S. ussuriensis A. Pojark 292 

9. S. elegans A. Pojark 293 

10. S. media Schmidt 294 

11. S. sericea Maxim 297 

12. S. dahurica Maxim 297 

13. S. alpina Pall 298 

14. S. tianschanica A. Pojark 299 

15. S. trilobata L 299 

16. S. pilosa Franch 300 

17. S. pubescens Turcz 301 

18. S. crenata L 301 

19. S. lasiocarpa Kar. et Kir 302 

20. S. ferganensis A. Pojark 303 

21. S. hypericifolia L 303 

22. S. aquilegifolia Pall 305 

Genus 719. Sibiraea Maxim. 

1. S. altaiensis (Laxm.) C. K. Schn. . 306 

2. S. tianschanica (Krassn.) A. Pojark 309 

Genus 720. Aruncus (L.) Adans. 

1. A. vulgaris Raf 310 

2. A. asiaticus A. Pojark 311 

3. A. kamtschatieus Rydb 311 

4. A. parvulus Kom 312 

Genus 721. Sorbaria A; Br. 

1. S. Olgae Zinserl 313 

2. S. sorbifolia (L.) A. Br 313 

3. S. Pallasii (G. Don) A. Pojark 315 

4. S. rhoifolia Kom 315 

Genus 722. Spiraeanthus Maxim. 

1. S. Schrenkianus (Fisch. et Mey.) Maxim 316 

Genus 723. Exochorda Li dl. 

1. E. Albert! Rgl 317 

2. E. tianschanica Gontsch 317 

Subfamily 2. Pomoideae Focke 

Genus 724. Cotoneaster Med. 
Section 1. Orthopetalum Koehne 

1. C. melanocarpa A. Pojark 320 

2. C. lacida Schlecht 323 

3. C. integerrima Medik 323 

4. C. uniflora Bge 324 

5. C. oligantha A. Pojark 326 

Section 2. Chaenopetalum Koehne 

6. C. multiflora Bge 329 

7. C. insignis A. Pojark 330 

8. C. racemiflora (Desf.) C. Koch 331 



9. C. taurica A. Pojark 332 

10. C. saxatilis A. Pojark 333 

Genus 725. Cydonia Mill. 

1. C. oblonga Mill 334 

Genus 726. Pyrus L. 

1. P. communis L 333 

2. P. Balansae Decaisne 339 

3. P. turcomanica Maleev \ 340 

4. P. Boissieriana Buhse 341 

5. P. ussuriensis Maxim 341 

6. P. asiae-mediae M. Popov 342 

7. P. Grossheimii A. Feck 345 

8. P. elaeagrifolia Pallas 346 

9. P. taochia G. Woron 347 

10. P. salicifolia Pall 348 

11. P. Takhtadzhiani A. Fed 349 

12. P. syriaca Boiss 350 

13. P. Sosnovskii A. Fed 350 

14. P. oxyprion G. Woron 351 

15. P. Raddeana G. Woron 351 

16. P. zang-ezura Maleev 352 

17. P. Korshinskyi Litw 352 

18. P. Regelii Rehd 355 

Genus 727. Ma/us Mill. 

1. M. silvestris Mill 359 

2. M. praecox (Pall.) Borkh 360 

3. M. orientalis Uglitzkich 362 

4. M. Sieversii (Ldb.) M. Roem 363 

5. M. Niedzwetzkyana Dieck 364 

6. M. turkmenorum Juz. et M. Pop 364 

7. M. domestica Borkh 365 

8. M. prunifolia (Willd.) Borkh 366 

9. M. baccata (L.) Borkh 369 

10. M. Pallasiana Juz 370 

11. M. manshurica (Maxim.) Kom 371 

12. M. sachalinensis Juz 372 

Genus 728. Sorbus L. 

Subgenus 1. Eu-Sorbus Kom. 

1. S. domestica L 374 

2. S. sambucifolia Roem 375 

3. S. Schneideriana Koehne 376 

4. S. aucuparia L 376 

5. S. glabrata Hedl 377 

6. S. polaris Koehne 378 

7. S. sibirica Hedl 378 

8. S. amurensis Koehne 381 

9. S. Boissieri C. K. Schn 382 



10. S. commixta Hedl 382 

11. S. kamtschatcensis Kom 383 

12. S. anadyrensis Kom 383 

13. S. tianschanica Ruprecht 384 

Subgenus 2. Hahnia Medik. 
Section 1. Aria DC 

14. S. subfusca (Ldb.) Boiss 390 

15. S. Albovii Zinserl 390 

16. S. subtomentosa Zinserl 391 

17. S. colchica Zinserl 391 

18. S. velutina C. K. Schn " 391 

19. S. Buschiana Zinserl 392 

20. S. schemachensis Zinserl 392 

21. S. caucasica Zinserl • 395 

22. S. Woronowii Zinserl 395 

— S. scandica Fr 396 

23. S. armeniaca HedL • 396 

— S. aria Crantz 397 

24. S. Kusnetzovii Zinserl 397 

25. S. graeca (Spach) Hedl 397 

26. S. migarica Zinzerl 398 

27. S. Baldacei Deg. et Fritsch 398 

28. S. turcica Zinserl . 399 

29. S. taurica Zinserl 400 

30. S. obtusidentata Zinserl 400 

31. S. persica Hedl 401 

32. S. turkestanica (Franch.) Hedl 401 

33. S. dualis Zinserl 402 

Section 2. Torminaria DC 

34. S. torminalis Crantz 405 

P04 729. Micromeles Decne. 

1. M. alnifolia Sieb. et Zucc 406 

Genus * Eriobotrya Ldb. 

1. E. japonica Ldb 407 

Genus 730. Amelanchier Med. 

1. A. rotundifolia (Lam.) Dum 408 

2. A. integrifolia Boiss. et Hoh 409 

3. A. turkestanica Litw 410 

— A. canadensis (L.) Med 410 

— A. spicata (Lam.) C. Koch 413 

Genus 731. Pyracantha M. Roem. 

1. P. coccinea M. Roem 414 

Genus 732. Mespilus L. 

1. M. germanica L 414 

Genus 733. Crataegus L. 
Section 1. Pinnatifidae Zbl. 

1. C. pinnatifida Zbl 421 



Section 2. Sanguineae Zbl. 

2. C sanguinea Pall 422 

3. C. dahurica Koehne 423 

4. C. altaica Bge 424 

5. C. Maximowiczii C. K. Schn 428 

6. C. remotilobata H. Raik 428 

7. C. chlorosarca Maxim 429 

Section 3. Pentagynae Zbl. 

8. C. pentagyna Waldst. et Kit 430 

9. C. pseudomelanocarpa M. Pop 432 

Section 4. Orientales Zbl. 

10. C. orientalis Pall 433 

11. C. Szovitsii A. Pojark 434 

12. C. pontica C. Koch 435 

Section 5. Oxyacantha Zbl. 

— C. oxyacantha L 437 

13. C. Meyeri A. Pojark 438 

14. C eriantha A. Pojark 439 

15. C. taurica A. Pojark 440 

16. C. ucrainica A. Pojark 441 

17. C. sphaenophylla A. Pojark 441 

18. C. ambigua C. A. M 443 

19. C. volgensis A. Pojark 444 

20. C. transcaspica A. Pojark 447 

21. C. caucasica C. Koch . . ., 447 

22. C. atrosanguinea A. Pojark 448 

23. C. songorica C. Koch 449 

24. C. kyrtostyla Fingerhut 450 

25. C. turkestanica A. Pojark 451 

26. C. Stevenii A. Pojark 453 

27. C. Beckeriana A. Pojark 453 

28. C monogyna Jacq 454 

29. C. pseudoheterophylla A. Pojark 456 

30. C. turcomanica A. Pojark 457 

31. C. microphylla C. Koch 459 

32. C. tianschanica A. Pojark 461 

33. C. dshungarica Zbl 461 

34. C. dipyrena A. Pojark 462 

35. C. zangezura A. Pojark 463 

36. C. Schraderiana Ldb 464 

37. C. pseudoazarolus M. Pop 465 

38. C. pseudoambigua A. Pojark 466 

39. C. armena A. Pojark • 467 



PREFACE 

The present volume, No. IX of the "Flora of the USSR," covers the families 
Droseraceae, Crassulaceae, Saxifragaceae, Hamamelidaceae, and Platanaceae, 
as well as the subfamilies Spiraeoideae and Pomoideae of the Rosaceae. 
Other subfamilies of Rosaceae are dealt with in Vol. X, as they could not 
possibly be accommodated in the present book. 

The present volume includes such plants of economic importance as 
currants and gooseberries, pears, apples, and related genera. The wide 
range of species of these genera growing wild in the USSR are of interest 
to breeders as material for remote hybridization, selection of wildlings, 
and direct introduction. This material could be of great value for 
orchards and soft-fruit plantations in the USSR. 

Many rosaceous species described in this volume have yielded, or may 
yield in the future, a considerable number of trees and shrubs for 
horticultural use. Numerous species of the Saxifragaceae and Crassulaceae 
may likewise prove to be of interest as ornamental plants. 

The study of the material treated in this volume has led to the identification 
of about 70 new species. In conformity with international rules, the 
descriptions pertaining to these species are also given in Latin. 

It should be noted that this volume brings the number of species up to 
about half the total number of higher plants of the USSR, which presumably 
does not exceed 14,000 species. 

The Editors 



Order 22. Sar r ac en ial e s engl. 

Flowers spirocyclic or cyclic, the perianth uniseriate or differentiated' 
into calyx and corolla; stamens inserted at the base of pistil; ovary 
superior; carpels 3—5; ovules numerous; seeds small, with endosperm; 
archesporium unicellular.— Herbs; leaves spiral, sometimes verticillate, 
more often arranged in basal rosette, entire, adapted for trapping insects. 



Family LXXII. DROSERACEAE DC. 

Perennial marshy or aquatic plants adapted for trapping and digesting 
insects; flowers regular, pentamerous, sepals and petals 5; stamens 5, 
anthers extrorse; pistil 1; styles 3— 5, simple or bifid; ovary unilocular, 
with 3—5 parietal placentae; fruit a capsule dehiscing by 3—5 valves; seeds 
small, mostly numerous. 

1. Leaves in whorls at the nodes of the filiform stem; aquatic plants . . . 

Genus 693. Aldrovanda Monti. 

+ All leaves rosulate, covered by reddish glandular hairs; plants of 

mossy, mostly sphagnum bogs Genus 694. Drosera L. 



Genus 6 93. ALDROVANDA* MONTI 

Monti Comment, bonon. acad.11,3 (1747) 404, tab. 12; L. Nov. plant . gen. ( 1751) 39. 

Aquatic plants with filiform stem; petioles broad, flat, cuneate, terminating 
in bristles, leaf blade of 2 semicircular lobes folding up along midrib; 
pedicels axillary, 1 -flowered; styles 5; stamens with cordate anthers; 
capsule globose, dehiscing by 5 valves; seeds ovaloid, black, smooth, 
shining. 

1. A. vesiculosa L., Sp.pl. (1753); 281; Ldb., Fl. Ross. I, 262; Shmal'g., 
Fl. I (1895) 118; N. Busch, Fl. cauc. crit. Ill, 4 (l 910) 744. - Ic. : Engl. 
Pflzr.IV, 112 (1906) f. 20; Kom. and Alis., Opred. rast. Dal'nevost. kraya I, 
Plate 178.- Exs.: Fl. pol. exs. no. 236. 

Perennial; stem somewhat branching 5—10 cm long; leaves in whorls of 
6—9, connate at base; petioles 0.5— 0.9 cm long, terminating in 6 subulate 
bristles; leaf blade rounded, 0.5— 0.7 cm long, 0.8— 1 cm broad; sepals elliptic 



After Ulysses Aldrovandi, professor of botany in Bologna (1522-1605). 



(3) 




PLATE I. 1-Aldrovanda vesiculosa L.: a) leaf, b) flower, c) pistil; 2-Drosera rotund i' 
folia L.: a) pistil and stamen, b) capsule, c) seed; 3-D.anglica Huds.: a) pistil and stamen. — 
4. D.obovata Mert.et Kolch.- D. intermedia Hayne: a) pistil and stamen, b) seed. 



or oval-elliptic, hairy on the margin; corolla white; petals obovate, 
0.4—0.5 cm long, ca. 0.25 cm broad. (Plate I, Figure 1 ). 

Distribution sporadic; known only from a few localities; most 
often floating to the surface in small oxbow lakes. European part; 
V.-Don (Khrenovskoi Bor), M. Dnp., Bl., L. V. (delta of the Volga; Caucasus: 
Cisc. (delta of the Kuban River), W. Transc. (Lake Bebesyr. Far East: 
Uss., Ze.-Bu.; Central Asia: Kyz. K. (delta of the Amu Darya). 
Gen. distr.: Scand., Centr. and Atl. Eur., W. Med., Centr. Afr., Bal. -As. Min., 
Jap.-Ch. (japan), Aust., E. India. Described from W. Europe. Type in 
London. 



Genus 6 94. DROSERA L. 

L.,Gen.ed. 1 (1737) 253. 

Calyx deeply 5-lob'ed; styles 3—5, deeply bifid; fruit a capsule dehiscing 
by 3—5 valves and enveloped by the persistent calyx and corolla; seeds 
numerous. Perennial marsh plants with a basal leaf-rosette; the upper 
surface of leaves covered with glandular hairs which secrete a digestive 
liquid acting upon the trapped insects. 

1. Capsule longitudinally grooved; stems usually arched -upcurved at 
base, ascending 3. D. intermedia Hayne. 

+ Capsule not grooved, more or less smooth; stems erect 2. 

2. Leaves prostrate with rounded blade 1. D. rotundifolia L. 

+ Leaves usually upright, linear -cuneate or obovate 3. 

3. Leaf blade linear-cuneate; capsule well developed ... 2. D. anglica Huds. 
+ Leaf blade obovate; capsule usually poorly developed 

4. D. obovata Mert. et Kock. 

1. D. rotundifolia L., Sp.pl. (175 3) 282; Ldb., Fl. Ross. I, 261 ; Shmal'g., 
Fl. I, 117; Turcz., Fl.baic.-dah. I, 191; Kryl., FL Zap. Sib. VI, 1397; Kom., 
Fl.Kamch. II, 196.-Rorella rotundifolia All. Fl. Pedem. II (1785) 88.- 
Ic. : Rchb., Ic. Fl. Germ. Ill, tab. XXIV, f. 4522; Dreves et Hayne, Bot. 
Bilderbuch (1798) tab. 2; Fl. Yugo-vost. V (l 931)467.- Exs.: Fl. pol. exs. 
No. 121; Meinsh., Herb. Fl. Ingr., No. 83; Herb. Fl. Rep. Sov. Ucr., No. 67; 
PL Finl. exs., No. 697, 698. 

Perennial; leaves prostrate, suborbicular to oblately rounded-oval, 
0.4—1 cm long, 0.45— 1.8 cm broad; capitate glandular hairs on the upper 
surface of leaves 0.5—1 mm long at center, 4— 5 mm at margin; petioles 
glabrous, 1— 7 cm long; scapes 1—3,7— 25 cm long, overtopping the leaves; 
flowers white, small, in rather loose terminal, racemes; calyx tubular - 
campanulate, 3.5—4 cm long, elongating in fruit to 5.5 mm long, the lobes 
oblong-obovate, obtuse; petals slightly exceeding calyx, oblong -obovate; 
capsule elongate -ovaloid, smooth, 5—6.5 mm long, 2—2.5 mm broad; seeds 
fusiform to 1.5 mm long, grayish, almost smooth. June — August. (Plate I, 
Figure 2). 

Sphagnum bogs and wet sand.— European part: all regions except 
BL, L. V., and, usually, L. Don; Caucasus (rarely): Cisc. and W. Transc; 
W. Siberia: Ob., U. Tob., Irt., Alt.; E. Siberia: all regions, but very rare 
in Dau.; Far East: Ze. -Bu., Kamch., Uda, Sakh.; Gen.distr.: Scand., 
Centr. and Atl. Eur., Jap.-Ch. (japan), N. Am. Described from W. Europe. 
Type in London. 



Economic importance. Medicinal; in the Shenkursk District and in the 
Vologda Region peasant women steam Drosera leaves in earthenware 
jugs: the leaves produce a proteolytic enzyme which is active even in 
the pores of the jug. 

2.. D. anglica Huds. .Fl.Angl.ed.il, 1 (1778) 35; Shmal'g., Fl. I, 117; 
Kryl., Fl. Zap. Sib. VI, 1398; Kom., Fl. Kamch. II, 198. - D. 1 o ng i f o 1 i a L., 
Sp. pi. ed.l (1753) 282 pp.; Ldb., Fl. Ross. I, 261 ; Turcz., Fl. baic. -dah. I, 
191.- R ore 11a longi folia Gilib., Fl. lith. V (1798) 141.- Ic: Rchb., 
Ic. Fl. Germ. Ill, tab. XXIV, f. 4524; Dreves et Hayne, Bilderb. (1798) tab. 3, 
f. A; Engl., Plfzr. IV, 112 (1906)96; Fl. Yugo-Vost. V (l 931 ) 467, fig. 407. - 
Exs. : Herb. Fl. Rep. Sov. Ucr., No. 68; PI. Finl. exs., No. 699; Fl. Ingr. exs., 
No. 84. 

Perennial; leaves obliquely ascending, linear -oblong or linear -cuneate, 
rounded at the apex, gradually tapering toward base, 1.5—4 cm long, 
0.2— 0.5 cm broad; petioles glabrous, 2— 9 cm long; scapes 1—3, 10— 25 cm 
long, surpassing the leaves; flowers somewhat larger than in preceding 
species; calyx to 0.5 cm long; petals obovate, 5—6 mm long, 3.5—4 mm 
broad; seeds ca.l mm, dark, oblong-fusiform. Other characters as in 
the preceding species. July — August. (Plate I, Figure 3). 

Peat bogs, mostly sphagnum.— European part: all regions except Bl., 
L. Don, and L. V.; W.Siberia: all regions; E.Siberia: all regions, but 
very rare in Dau.; Far East: Ze.-Bu., Uss., Uda, Kamch., Sakh. 
Gen. distr. : Scand., Centr. and Atl. Eur., Bal. -As. Min. (rarely), Mong. 
(Tuva), N.Am. Described from England. Type in London. 

3. D. intermedia Hayne, Bot. Bilderbuch (1798) 18; Ldb., Fl. Ross. I, 261; 
Shmal'g., Fl. I, 117; N. Busch, Fl. cauc. crit. Ill, 4 (1910) 745.- D. longifolia 
L., Sp.pl. (1753) 282 p. p.- Ic: Engl., Plfzr. (D r o s e r a c e a e), IV, 1 12 (1906) 
95.- Exs.: HFR, No. 3569 and 2009; Herb. Fl. Rep. Sov. Ucr., No. 66 ; PL 
Finl. exs., No. 248a, 248b, 248c. 

Perennial; leaves in radical rosettes, cuneate -obovate, erect, the blade 
7— 12 mm long and 2—4 (4.5) mm broad; petioles 1.3— 3 cm long; stems 
usually 1—3, arched -upcurved at base and ascending, 5—8 cm long, usually 
only slightly surpassing the leaves; sepals ovate, obtuse, appressed, reflexed 
at summit; petals white; capsule pyriform, with 3—4 longitudinal grooves: 
seeds tuberculate, with adherent testa. July— August. (Plate 1, Figure 5). 

Peat bogs, notably moorland.— European part: Lad.-Ilm., especially in 
coastal regions (Lakhta, Sestroretsk, Lisii Nos, and near Pskov), U. Dnp.; 
Gen. distr. : Scand., Centr. and Atl. Eur., N. Am., W. Indies (Cuba). Described 
from Europe. 

4. D.obovata Mert. et Koch, Fl. Deutschl. II (1826) 502.- D. rotund i - 
folia X anglica Lasch in Bot. Zeit. XV (1857) 514; Shmal'g., Fl. I, 1 17; 
Kom., Fl. Kamch. II, 197. 

Perennial; leaves mostly erect, oboval, obovate, or spatulate, 1—2.2 cm 
long, 0.4— 0.8 cm broad; petioles 4— 5 cm long; stem 10— 15 (20) cm high, 
erect, glabrous; raceme 1—8 -flowered; stigmas 2 -lobed; capsule often 
reduced, shorter than calyx; seeds obovoid, ca. 1 mm long. (Plate I, 
Figure 4). 



Distribution as for D. r ot u nd i fo 1 ia and D. a ng 1 i c a, and probably a 
hybrid between the two. A hybrid D. rotundifoliaX D. intermedia 
has also been described (Callier, Schrift. Schles. Ges. II, 1892-1894). 



Order 23. RosaleS LINDL. 

Flowers bisexual, rarely unisexual; regular to zygomorphic, mostly 
pentamerous, cyclic or rarely spirocyclic; perianth mostly biseriate, 
petals rarely rudimentary; stamens as many or twice as many as 
petals, or indefinite, rarely 1—4; gynoecium superior or inferior; carpels 
as many as petals or fewer, rarely numerous, free or more or less united; 
placentation marginal; pollen grains binuclear; archesporium usually 
multicellular. Plants of widely differing aspect; leaves with or without 
stipules. 

1. Fruit a pod; flowers usually zygomorphic, papilionaceous, less often 

nonpapilionaceous and then regular or irregular 

Family LXXVIII. Leguminosae Juss. 

+ Fruit different; flowers usually actinomorphic 2. 

2. Herbaceous plants, sometimes undershrubs 3. 

+ Trees or shrubs 6. 

3. Leaves exstipulate 4. 

+ Leaves stipulate Subfamily Rosoideae (family Rosaceae). 

4. Carpels as many as petals, i. e., mostly 5, rarely 3—4 by abortion ... 5.. 
■+ Carpels fewer than petals, mostly 2 

Family LXXIV. Saxifragaceae DC. 

5. Leaves simple, often fleshy Family LXXIII. Crassulaceae DC. 

+ Leaves pinnately compound, herbaceous 

Genus Aruncus (family Rosaceae). 

6. Flowers unisexual 7. 

+ Flowers bisexual (in Soviet genera) 8. 

7. Flowers in loose clusters; leaves entire 

Family *Eucommiaceae Van-Tiegh. 

+ Flowers in compact capitate inflorescences; leaves mostly lobed . . . 

Family LXXVI. Platanaceae Lindl. 

8. Flowers in groups of 2—5, forming axillary capitate inflorescences 
Family LXXV. Hamamelidaceae R. Br. 

+ Flowers differently arranged 9. 

9. Plants with evergreen leaves and small fragrant flowers; fruit a 
capsule Family *Pittosporaceae Lindl. 

+ Plants with a different combination of characters 10. 

10. Fruit composed of achenes, sometimes borne at the bottom of an 
enlarged hypanthium, forming a spurious fruit, or else fruit an 
aggregate Subfamily Rosoideae (Family Rosaceae Juss.). 

+ Fruit not as above 11. 

11. Leaves opposite, exstipulate; ovary inferior or half-inferior, 

2— 5-locular; fruit a capsule dehiscent along the sutures 

Subfamily Hydrangeoideae (Family Saxifragaceae DC.). 



+ Leaves alternate 12. 

12. Fruit a juicy many -seeded berry; ovary inferior, styles 2, mostly 
connate; flowers in racemes (these sometimes reduced to 1—3 
flowers) Subfamily Ribesioideae (family Saxifragaceae). 

+ Fruit an aggregate of follicles, a drupe, or a pome 13. 

13. Carpels usually 5; fruit an aggregate of 5 (2—7) free or more or less 
(sometimes completely) united follicles, dehiscing along the inner 

suture; flowers small, in corymbs, panicles, or spikes 

Subfamily Spiraeoideae (family Rosaceae Juss.). 

+ Carpels 1—5; fruit a drupe or pome; flowers in corymbs or 

umbels 14. 

14. Carpel 1, not united with hypanthium; fruit a drupe 

Subfamily Prunoideae (family Rosaceae Juss.). 

+ Carpels 1—5, united with inner side of hypanthium; fruit a pome 

Subfamily Pomoideae (family Rosaceae Juss.). 



Family LXXIII. CRASSULACEAE DC* ^ 

Flowers actinomorphic, bisexual or unisexual, the plants then usually 
dioecious, arranged in umbels, spikes, or panicles, sometimes solitary; 
sepals 3—20, almost free ©r partly connate; petals as many as calyx-lobes, 
free or connate, adnate to stamens, sometimes inconspicuous or lacking 
(Penthoroideae); stamens as many or twice as many as petals, free or 
adnate to petals; carpels as many as petals, with nectariferous scales at 
base; fruit comprising membranous or leathery follicles free or connate 
at base, or united nearly to the middle (Penthoroideae), dehiscing along 
the ventral suture or by a ring of transverse clefts (Penthoroideae); seeds 
numerous, small. Herbaceous annuals or perennials, rarely undershrubs 
with a woody stock; leaves mostly succulent, fleshy, simple, exstipulate. 

Note. The family Crassulaceae numbers ca. 1,500 species in 6 
subfamilies and 33 genera distributed all over the globe. About 550 species 
in 10 genera belong to the subfamily Sedoideae. This subfamily includes 
most of the species of Crassulaceae occurring in the USSR (107). The 
Semperviroideae with ca. 85 species in 4 genera are distributed throughout 
the Mediterranean region, the Canary and Madeira Islands. All the 
species growing in the USSR belong to Sempervivum. The subfamily 
Cotyledonoideae, with over 70 species in 6 genera, is represented in the 
Soviet Union by a single species of Umbilicus (subgenus Chiasto - 
phyllum) growing in the Caucasian forests. The Penthoroideae, which 
could well be considered as a distinct family, consists of the single genus 
Penthorum, which is distributed in the Soviet Union and in North America. 
The Crassuloideae, with ca. 5 genera and 300 species in tropical Africa, 
S. Arabia, and Madagascar, are represented in the USSR only by T i 1 1 a e a. 
The subfamilies Kalanchoideae (ca. 230 species) and the Echeverioideae 
(ca. 200 species) do not occur in the USSR. 

Economic importance. Owing to the variety of forms and the relative 
ease of vegetative propagation, the Crassulaceae deserve special attention 
as ornamental plants suited to widely different climatic zones. Numerous 

* Treatment by A.G. Borisov. 



species are widely cultivated, such as certain species ofSempervivum 
and Sedum (S. sparium, S. spectabile, and many others). The cell 
sap of the Crassulaceae contains malic acid, traces of tartaric acid, 
and tannin (Hegi). 

Note. As they become much deformed when dry, it is essential to 
study the Crassulaceae in viva. 



Key to Genera 

1. Fruits free or connate at the base, dehiscing along ventral suture; 
flowers 3— 20-merous, always with conspicuous corolla and calyx. 
Mostly succulent plants 2. 

+ Fruits connate nearly to the middle, dehiscing by a ring of transverse 
clefts; flowers 5 (6— 8)-merous, petals inconspicuous or lacking. Plants 

not succulent.— Far East. (Subfamily 5. Penthoroideae Engler) 

Genus 703. Penthorum L. 

2. Stamens as many as petals; flowers 3—4 -merous, petals distinct; 
leaves opposite; small annual plants growing in moist sandy -silty 

soils. (Subfamily 1. Crassuloideae Berger) 

. Genus 695. Tillaea (Mich.). 

+ Stamens twice as many as petals, or of same number as petals but 
then leaves alternate; flowers 4— 20-merous; petals distinct or 
connate 3. 

3. Calyx and corolla 6— 20-merous; stamens twice as many as petals; 
petals distinct; calyx cup-shaped; leaves fleshy, alternate, the radical 

rosulate. (Subfamily 3. Sempervivoideae Berger) 

Genus 697. Sempervivum L. 

+ Plants not as above 4. 

4. Calyx and corolla 5 -merous; stamens twice this number; petals connate 
nearly to the middle; leaves opposite, decussate, flat, petioled; basal 

rosettes lacking; plants with creeping rhizome 

Genus 696. Umbilicus C. C. (subgenus Chiastophyllum Stapf.). 

+ Calyx and corolla 4—6 (9)-merous ; stamens as many or twice as many 
as petals; petals distinct or connate and then leaves alternate; leaves 
of various shape, alternate, opposite, whorled, or rosulate; underground 
parts various, if a creeping rhizome then petals distinct. (Subfamily 4. 
Sedoidea Berger) 5. 

5. Plants with basal leaf rosette 6. 

+ Plants without basal leaf rosette 7. 

6. Rosette leaves usually cuspidate from a white cartilaginous apical 
appendage, rarely soft -acuminate or subobtuse; flowering shoots 
solitary from center of rosette; flowers 5 -merous, in compact spiciform 
racemes; petals distinct, spreading. S. Siberia, from the Urals to China and 

the Far East, to the south as far as T. Sh. and Pam. -Al 

Genus 701. Orostachys Fisch. 

+ Rosette leaves muticous, without cartilaginous appendage; flowering 
shoots usually numerous, from axils of basal leaves, rarely solitary 
from center of rosette; flowers 5—6 (7)-merous, loose, in corymbs or 



11 



corymbiform panicles; petals campanulately connate, the corolla lobes 

erect. Centr. Asia, to Altai in E., to As. Min. in W 

Genus 702. Rosularia Stapf. 

7. Plants with a sturdy, woody, vertical, sometimes branching condex cov 
covered by crowded, appressed, chaffy leaves, and simple stems arising 
from their axils; flowers small, 4— 5-merous, dioecious, sometimes 
bisexual, the petals distinct; inflorescences corymbiform to capitate- 
corymbiform with yellow or red flowers, or spiciform racemes with 
white or pink flowers. Subgenus Clement sia (Rose) A. Bor.). 
Arcto-montane species . Genus 698. Rhodiola L. 

+ Plants with tuberous roots or creeping rhizome, or with shortened 
rhizome and a tuft of thin roots, or annual plant with thin roots; 
underground parts without scalelike leaves or, if few such leaves 
present, petals connate to above middle, forming a large infundibular 
or campanulate corolla; flowers (4) 5— 9-merous, always bisexual ... 8. 

8. Flowers 5 - or 6-merous; corolla infundibular or campanulate, with 
straight lobes, pinkish or reddish, drying golden yellow; petals 
connate to the middle; leaves fleshy, oblong to linear, glabrous, 
perennial; herbs with erect stems, the short caudex sometimes covered 

with small scalelike leaves. Central Asia 

Genus 700. Pseudosedum Berger. 

+ Flowers usually 5-merous, more rarely (4) 6— 9-merous, corolla usually 
with stellately recurved lobes, yellow, white, pink, or red; petals 
distinct or basally connate. Annual and perennial herbs of various 
aspect Genus 699. Sedum L. 



Subfamily 1. CRASSULOIDEAE Berger in Engler. u. Pr. Nat. Pflzfm. 
18a (1930) 386.— Stamens in 1 series, as many as petals; flowers (in Soviet 
plants) 3- or 4-merous; petals usually absent; leaves opposite; small 
annual plants. 



Genus 6 95. TILLAEA L.* 

L.Gen.pl.ed.5 (1754) 62. 

Sepals and petals 3 or 4; stamens 3 or 4; pistils 3 or 4; follicles many- 
seeded. Small annuals, 2— 5 cm high, with cylindrical opposite leaves; 
flowers small, terminal, or in the leaf axils or rarely in the forks of stem. 
Plants of rather moist sandy and clayey habitats. 

1. Flowers sessile or borne on very short pedicels; stem internodes 
shorter than leaves; leaves dense, linear-oblong or oblong-lanceolate, 
acuminate 2. 

+ Flowers on pedicels 2—2.5 times as long as the leaves; internodes 

longer than leaves; leaves few, linear, obtuse ... 1. T. vaillantii Willd. 

2. Flowers sessile, singly in upper leaf axils; stem internodes distinct 
2 . T. aquatica L. 

* Named for the Italian botanist Michelangelo Tilli ( 1663-1740), author of a catalog of plants of the Pisa 
Botanical Gardens. 



10 



+ Flowers borne on very short pedicels, one in the axil of each of a 
pair of decussate leaves from the base of the stem, the crowding of 
leaves resulting in a spiciform inflorescence; internodes indistinct 

3. T. alata Viv. 

1. T.vaillantii Willd., Spec. pi. I (1798) 720; Shmal'g., Fl. I, 359. - 
T.aquatica Lam., Illustr. I (1791 ) 361, non L. - T. s a g i no id e s Maxim. 
in Bull. Ac. Petersb. XXVI (1880)473.- Bulliar da vaillantii DC. 
PI. gras. (1801) tab. 74.- B.aquat ica Fedtsch. in Consp., Fl. Turk. 3 
(1909) 68, non DC- Crassula vaillantii Schoenl. in Engl, und Prantl, 
Nat. Pflzfm. Ill, 2a (1891) 37; Berger in Engl., Nat. Pflzfm. 18a (1930) 388.- 
Tillaeastrum vaillantii Britton in Bull. N. Y. Bot. Gard. Ill (1903) 2. - 
Ic: Borisova in Fl. Yugo-Vost.,No. V (1931) f.408; Rchb., Ic. Fl. Germ. 
XXIII (1898-1899) tab. 40; Lam., 1. c, tab. 90; DC., 1. c, tab. 74. 

Biennial, 2-5 cm high; stem simple or branching, erect or prostrate, 
the few -leaved internodes long; leaves decussate, remote, linear, obtuse, 
glabrous, sheathing at base, 1.5-3 mm long, ca. 0.5 mm broad; flowers 
solitary in axils of leaves, sometimes in stem forks; pedicels 4-6 mm 
long, 2-2.5 times length of leaves; flowers 4-merous, small, ca. 1.5-2 mm 
long; calyx glabrous, basally adnate to corolla, lobed to the middle; the 
lobes broadly triangular, obtuse; petals pink, ovate, twice as long as calyx; 
stamens 4, exceeding calyx but shorter than petals with rounded anthers 
and short filaments; hypogynous scales linear, alternating with stamens; 
follicles usually 6-8-seeded, ovaloid, short -acuminate; seeds oblong- 
ovaloid, dark brown, ca. 0.4 mm long, 0.1-0.2 mm broad. May -June. 
(Plate II, Figure la, b). 

Boggy sites, compact solonetzic soil, less often riverbanks and 
lakeshores.- European part: Bl. (between Voznesensk and Pervomaisk), 
L. V. (Krasnoarmeisk, Chapchachi); Central Asia: Ar. -Casp. (Air-tau; 
between Lakes Sary-kul' and Dzhumor-kul', Lake Sassyk-kul'), Balkh. 
(banks of Black Irtysh River). Gen. distr. : Med., N. Am. Described from 
southern France. Type in Berlin. 

2. T.aquatica L., Sp.pl. (1753) 12 8; Shmal'g., Fl. I, 359. - T. pro st rat a 
Schkuhr in Usteri Ann. Bot. XII (1794) 6.- T. simplex NuttaL, in Journ. 
Acad.Phil. I (1817) 114.- Bulliar da aquatic a DC. in Bull. Soc. Philom., 
No. 49 (1801)2; Prodr. HI, 382; Ldb., Fl. Ross. II, 1 72. - B. p r o s t r at a 
Weinm., Fl. petrop. (1837) 21. - B. s c hkuhr i i Spreng., Syst. I (1825) 498.- 
Tillaeastrum aquaticum Britton in Bull. N. Y. Bot. Gard. Ill (l 903) 1 . - 
Crassula a qu at ica Schoenl. in Engl. u. Prantl., Nat. Pflzfm. Ill, 2a 
(1891) 37; Berger in Engl. u. Pr., Nat. Pflzfm. 18a (1930), 389.- Ic: Rchb., 
Ic. Fl. Germ. XXIII (1898-1899) tab. 39; Hegi, 111. Fl. Mitt. Eur. IV, 2, f. 902. - 
Exs.: HFR, No. 18, No. 2021; PL Finl. exs., No. 249. 

Annual or biennial, glabrous, 1-3 (5)cm high, with fibrous roots, densely 
leafy; stems branching, in water erect, in wet damp habitats usually 
prostrate and rooting at the nodes; leaves decussate, sheathing at base, 
linear, 4-6 mm long, 0.5-1 mm broad, thickish, entire, acute, glabrous; flowers 
small, 4-merous, subsessile, 1 or 2 in the leaf axils; calyx basally connate, 
with broadly triangular pointed lobes; petals whitish, ovate, 1—1.5 mm long, 
0.5— 0.8 mm broad, erect, 1.5-2 times as long as calyx; stamens 4, shorter 



11 



13 



than petals; hypogynous scales filiform, alternating with stamens; fruit 
of 4 follicles, ovaloid, ca. 1.3 mm long, 0.5 mm broad, terminating in a 
subulate beak; seeds numerous, 8—10, ovaloid, 0.3—0.6 mm long, 0.2 mm 
broad, dark brown, longitudinally rugose. June —September. (Plate II, 
Figure 2a— b). 

Infrequent, mostly along riverbanks, on sandy soils, silty and moist 
sites, seashores. European part : Kar.-Lap. (Svir River, Lake Onega), 
Dv.-Pech. (Arkhangelsk — reported by Schmalhausen), Lad.-Ilm., 
U. Dnp. (Minsk — reported by Schmalhausen); Far East: Uss. (surroundings 
of Olga and Milogradovo villages). Gen. distr. : Scand., Centr. Eur. (north 
as far as Iceland), N.Am. (California), Jap.-Ch. (japan, Korea). Described 
from Europe. Type in London. 

3. T.alata Viv. PI. Aeg. Dec. (1830) 16.- T.tr ichopoda Fenzl. in 
Ky. PL Pers. austr. exBoiss.,Fl. Or. II (1872) 767. - Crassula alat a 
Berger in Engl. u. Pr. Nat. Pflzfm. 18a (1930) 389. 

Annual, 1 — 3 cm high, often subcespitose; stems slender, ascending, 
simple or slightly branching, the internodes very short; leaves crowded 
along the stem, oblong-lanceolate to ovate -oblong, acuminate, sheathing 
at base; flowers ca. 1 mm long, subsessile, in axils of all leaves from the very 
base; calyx reddish, the distinct lobes oblong-ovate; petals shorter than 
sepals, whitish, oblong, filiform -acuminate; stamens 4, inserted at the 
base of petals, the filaments ! / 3 the length of calyx, the anthers rounded; 
follicles ovaloid, short -beaked, half the length of calyx; seeds ovaloid, 
dark brown, longitudinally grooved, subobtuse, 0.3 mm long, 0.2 mm broad. 
March. (Plate II, Figure 3a— b). 

Maritime sands, moist sandy sites.— Caucasus: Tal. (Ol'khovka village). 
Gen. distr. : E. Med. (Egypt, Syria — maritime part), Iran, (near Dalaki). 
Described from Egypt. 



Subfamily 2. COTYLEDONOIDEAE Berger in Engl. u. Pr. Nat. Pflzf. 18a 
(1930)412. Stamens in 2 series, twice as many as petals; flowers 
5-merous; petals connate nearly to the middle; leaves decussate and 
petioled, or alternate; basal rosettes absent. Plants with creeping 
rhizome or tuberiferous. 



Genus 6 96. UMBILICUS * DC. 

DCProdr.III (1838) 400, pro parte. 

Flowers 5-merous; sepals free, triangular, acute; corolla campanulate; 
petals connate nearly to the middle, with prominent midnerve, their 
lobes lanceolate -triangular, acute; stamens 10, shorter than corolla, the 
filaments adnate to throat of corolla; follicles free, linear -lanceolate, 
usually nutant at maturity; hypogynous scales linear. Glabrous plants 
with creeping rhizome (subgenus Chiastophyllum Stapf) or with 
tuberous, often rounded, underground part; leaves decussate, approximate 
toward base of stem, large, rounded -ovate, coarsely crenate-sinuate, 
abruptly narrowing to petiole, fleshy (subgenus Chiastophyllum) or 

* From the Latin word meaning navel, middle, center. 



5T73 12 



leaves alternate, peltate, or cordate, funnel-shaped at center, petioled 
(subgenus Eu -Umbilicus A. Bor.); flowers in racemes. 

Note. Species of Umbilicus (Eu-Umbilicus A. Bor.) occur 
in the Mediterranean region, eastward to Asia Minor; one species in 
Ethiopia. Only one species of the subgenus Chiastophyllum Stapf. 
occurs in the USSR. 



Subgenus CHIASTOPHYLLUM (Ldb.) Stapf. in Index Londin. II (1930) 176, 
316, pro genere; Berger in Engl., Nat. Pflzfm. 18a (l 930) 419; Sect. 
Chiastophyllum Ldb. Fl. Ross. II (1844-1846) 176; Boiss.Fl.Or.il, 
774.— Plants with creeping rhizome; leaves rounded -ovate, petioled, 
decussate. 

1. U. oppositifolius Ldb. Fl. Ross. II (1844-1846) 176; Boiss., Fl. Or. II, 
774; Grossh., Fl. Cauc. II, 231 . — Chiastophyllum o p p o s i t i f o 1 iu m 
Berger in Engl. u. Pr.,Nat.Pflzfm. 18a (1930) 419.- Cotyledon 
oppos itifolia Ldb. ex Nordmann in Bull. Acad. Pe'tersb. II (l 837) 313.— 
Sedum oppositifolium Raymond Hamet in Candollea IV (l 92 9— 1 931 ) 
43.- Ic: Bot. Mag. (1919) tab. 8822; Frod. in Acta Horti Gothoburg. VII 
(1932) pi. LVIII, textfigs 123-133. 

Perennial, 15—30 cm high, glabrous; rhizome creeping, long, with 
fibrous roots; stems decumbent; shoots densely leafy below, 
remotely above; flowering stems terminal, erect; leaves decussate, flat, 
the lower ones to 4 cm long, the upper smaller, rounded -ovate, petioled, 
obtuse, bluntly crenate -sinuate; inflorescence terminal, racemose-paniculate, 
many -flowered, erect, with shorter lateral racemes; racemes sometimes 
borne in the axils of 2 uppermost leaf pairs; bracts subulate, shorter than 
flowers, equaling the short pedicels; flowers alternate, remote, horizontal 
at anthesis, finally nodding; calyx 5 -parted, almost to the base; calyx lobes 
ovate, acuminate, appressed, as long as or slightly longer than corolla 
tube; corolla 4—5 mm long, creamy, 5 -cleft, almost to the middle; corolla 
lobes straight, pointed, ovate -lanceolate, lobes 2—2.5 times as long as the 
calyx; stamens 10, yellow like the anthers; filaments adnate to base of 
corolla; hypogynous scales 5, linear, slightly tapering toward base; 
follicles 5, linear-lanceolate, short -beakers, many-seeded; seeds small, 
0.5 mm or slightly longer, ovoid -lanceolate. Fl. May — June; fr. August. 
(Plate II, Figure 4a-b). 

At altitudes of 1,700—2,600 m, wooded slopes, beech forests, stony soils, 
spring banks, mainly calcareous regions, most frequent in W. Caucasus. — 
Caucasus: Cisc. (northern slopes of Main Range), W. Transc. (often), 
E. Transc. (rarely). Endemic. Described from Abkhazia. Type in Leningrad. 



Subfamily 3. SEMPERVIVOIDEAE Berger in Engl. u. Pr. Nat. Pflzfm. 18a 
(1930) 420.— Stamens in 2 series, twice as numerous as petals; flowers 
6—20 (32)-merous; petals distinct; calyx cup-shaped; leaves fleshy, convex, 
alternate, the basal rosulate. 



13 



15 



16 



Genus 697. SEMPERVIVUM L. 

L.Gen. pi. ed.5 (1754) 209. 

Flowers 6—20 -merous, white, yellow, yellowish green, red, pink, or 
purple; calyx green, fleshy, calyciform, basally connate; petals distinct, 
stamens 12—40 basally adnate to petals, carpels free; hypogynous scales 
small, entire or fimbriate at the apex. Fleshy, glabrous or glandular - 
pubescent perennials; leaves alternate, entire, fleshy, usually ovate or 
oblong, acute or acuminate, ciliate, on sterile shoots crowded into spherical 
basal rosettes, on flowering shoots remote and more elongated; flowering 
shoots arising from center of leaf rosettes; inflorescence a corymbiform 
panicle composed of scorpioid cymes. 

Approximately 25 species distributed throughout the mountains of central 
and southern Europe, to the Caucasus and the Volga in the east. Mainly on 
rocky, stony, and pebbly sites. 

Economic importance. Ornamental species of S e mp e rv ivum, known 
by over 200 names, are cultivated in various European gardens, forming 
numerous hybrids. The sap of S.tectorum contains a large amount of 
malic acid (Annenkov). Leaf rosettes and young shoots of S. s ob o 1 i f e r um, 
and of other species, are used as a salad vegetable. 

1. Flowers 6 -merous, campanulate; petals erect, fimbriate (Section 2, 
Jovisbarba DC.) 7. S. soboliferum Sims. 

+ Flowers 8— 18 -merous; petals stellate by spreading, entire. Section 1. 

Eu-Sempervivum Schoenl.) 2.j 

2. Flowers yellow or greenish yellow, often drying green 3... 

+ Flowers purple, red, pink, sometimes drying yellow 5. 

5. All leaves glabrous above, with long-ciliate margin; lobes mostly 

ovate, subobtuse; stamens about as long as petals 

6. S. glabrifolium A. Bor.i 

+ All leaves pubescent on upper surface; calyx lobes lanceolate or 

oblong-ovate, acute; petals 1.5 times as long as stamens 4., 

4. Inflorescence loose, many -flowered, 4—10 cm long, ca. 8—10 cm in 
diameter, with rather long scorpioid branches; corolla 3—4 times as 
long as calyx 4. S. ruthenicum (Koch) Schnittsp. et Lehm. , 

+ Inflorescence contracted, few-flowered, 1—3 cm long, 2— 5 (6) cm in 
diameter, with short branches; corolla 2—2.5 times as long as 
calyx 5. S. globiferum L. 

5. Rosette leaves glabrous or sparsely short -pubescent above, oblong - 
obovate, widest at the middle, short -acuminate, with soft or stiff 
hairs on the margin; cauline leaves more elongated to lanceolate; 
inflorescence many -flowered. Plants 12—60 cm high 6. 

+ Rosette leaves glandular -pubescent above, oblong-lanceolate, broadest 
at the base, long -acuminate margin soft pilose; cauline leaves 
lanceolate to linear -lanceolate; inflorescence few-flowered. Plants 
4—10 (20) cm high, grayish pubescent 1. S. pumilum M. B. 

6. Plants 12— 40 cm high; leaf rosettes 3— 5 cm in diameter; petals 

2—3 times as long as calyx; inflorescence 2—7 cm in diameter 

3. S. caucasicum Rupr. 

From the Latin semper, always, and vivu m, alive, referring to the viability of the very drought-resistant 
succulent leaves. 



14 



Plants 45—60 (lOO)cm high; leaf rosettes 5—7 (20)cm in diameter; 
inflorescence 7—10 (20)cm in diameter; petals up to 1.5 times as long 
as calyx 2. S. tectorum L. 



Section 1. EU-SEMPERVIVUM Schoenl. in Engl, und Prantl., Nat. Pflzfm. 
3, 11a (1890) 31; sect. J o v i b a r b a DC, PI. rar. jard. Gen. (1829), No. 21, 80, 
obs.; Prodr. Ill, 413 p. p.; sect. S e m p e r v i v u m genuinum C.Koch, 
Syn. Fl. Germ. (1837) 288.- Flowers 8-18-merous; petals stellately 
spreading, entire. 

1. S.pumilum M. B., Fl.taur.-cauc. I (1808) 381; Shmal'g., Fl. I, 363; 
Grossg., Fl.Kavk. II, 233; Berger in Engl. u. Pr., Nat. Pflzfm. 18a (1930) 
422; Praeger, An account Semperv. group (1932) 57.— ?S. montanum 
Eichw., Casp.-cauc. (1831-1833) 31, non L. - ? S. a Hum Turrill in Rec. 
prof. Petkoff (1936) 126.- Ic: Praeger, 1. c, f. 13. 

Perennial (2.5) 4— 10(20) cm high, white -puberulent; rosettes small, 
1—2.5 cm in diameter; their leaves oblong-lanceolate, abruptly short- 
acuminate, glandular -pubescent on both sides or glabrescent, with ciliate 
margin, green, 1 cm long, 3—4 broad, 2 mm thick, flat above, convex 
below; cauline leaves lanceolate to linear -lanceolate, acuminate, glandular - 
pubescent, 1 — 1.5 cm long; inflorescence 2 — 8 (lO)-flowered, with 2 or 3 
short, pubescent branches; flowers subsessile, 2—4 cm in diameter; 
corolla 3 times as long as calyx; calyx pilose, 5 mm long, basally connate, 
with 10—12 broadly lanceolate acute reddish lobes; petals 10—12, 
lance-linear, lilac -purple, with a darker median stripe, horizontally 
spreading, pilose on the outside and along the margins, with sparse hairs 
on the inside; stamens 20—24, 4—6 mm long, the purple filaments dilated 
toward the base; anthers red; hypogynous scales flat, oblong, obtuse, 1 mm 
long, green; fruit of 10 — 12 divergent follicles, 3.5—5 mm long, more or 
less glandular -pubescent, with reddish beak. Fl. July — August, fr. from 
August. 

Taluses, rocks, moraines near glaciers, mainly in the alpine zone, 
also among xerophytic mountain vegetation at 1 ,300— 3,000 m. — Caucasus: 
Cisc. and E. Transc. (often), Dag., S. Transc. (rarely), W. Transc. (rarely). 
Endemic. Described from the Caucasus (Terek River valley near 
Ordzhonikidze). Type in Leningrad. 

2. S. tectorum L., Sp.pl. (1753) 464; Ldb., Fl. Ross. II, 189, p. p.; DC, 
Prodr. Ill, 413; Berger in Engl. u. Pr.,Nat. Pflzfm. 18a (1930) 421.- 
Sedum tectorum Scop., Fl. Carn. II (l 772) 325. - Ic. : Rchb., Ic. Fl. 
Germ. XXIII, tab. 67; Hegi, 111. Fl. Mitt. Eur. IV, 2, tab. 141, f. 923, 924a-i; 
Praeger, An account Semp. group (1932) tab. 16. — Vernacular names: 
zhivuchka krovel'naya, skochki. 

Perennial, 45— 60 (lOO)cm high; stems glandular -pubescent with white 
hairs; rosettes (3) 5—7 (20)cm in diameter, open, flatfish; their leaves 
green, often becoming red, glabrous except ciliate margin, oblong-obovate, 
acuminate, 3—6 cm long, 1—1 .5 cm broad, fleshy, flat above, convex beneath, 
whitish at base; cauline leaves ovate -lanceolate, acute, glandular -pubescent, 
the margin densely long-ciliate; inflorescence dense, many -flowered, 



15 



profusely branching, 7— 10(20 ) cm in diameter, long -branched, flattened 
or obtriangular in outline; bracts linear, acute, pilose; flowers 12 — 16 — 
mostly 13-merous, to 2.5 cm in diameter; calyx ca. 8 mm long, pilose, divided 
to the middle into lanceolate, acute lobes; petals dark or light purple or 
red, greenish-nerved, stellately spreading, 9—12 cm long, glandular -pubescent 
on the back and margin, linear -lanceolate, acute; stamens 24—32 (mostly 26), 
shorter than petals, 5—6 mm long, with red filaments and anthers; 
hypogynous scales suborbicular, green; fruit of 12—16 follicles green, 
glandular-pilose, ca. 7 mm long, the subulate red glabrous beak half 
follicle length. Fl. July— September. 

Rocky and stony sites, often cultivated or escaped.— European part: 
recorded for U. Dnp. and M. Dnp. (near Zhitomir, Mogilev, and 
Dnepropetrovsk, according to Paczoski and Schmalhausen), probably 
erroneously, or having become naturalized as in Poland and the Baltic 
republics. Gen. distr. : Centr. Eur., Med., Bal. -As. Min. Described from 
Europe. Type in London. 

Economic importance. S.tectorum L. is widely cultivated as an 
ornamental plant and is extensively used for covering adobe and thatch 
roofs; hence its specific name. Medicinal plant. 

3. S. caucasicum Rupr. ex Boiss., Fl. Or. II (1872) 796.- Grossg., Fl. 
Kavk. II, 232; Praeger, An account Semp. group (1932) 76.— S.tectorum 
auct., non L. - S.montanum C. A.M., Verz. Pfl. Cauc. (1831) 152, 
nonL.; Ldb., Fl. Ross. II (l 844-1846) 189, non L. - S. t e ct o ru m subsp. 
caucasicum Berger in Engl. u. Pr.,Nat. Pflzfm. 18a (1930) 422.- ? 
S. f 1 ag e 1 1 i f e r um Fisch. in herb. 

Perennial, (10) 12— 20 cm high; stems glandular -pubescent with white 
hairs; rosettes 3—5 cm in diameter, their leaves green, oblong -obovate, 
acute, glabrous or puberulent, with stiffly ciliate margin; cauline leaves 
lanceolate, acute, 2—5 cm, glandular -pubescent, the margin long-ciliate; 
inflorescence many -flowered, branching, corymbiform, stoutly short - 
branched, 2—7 cm in diameter; bracts linear, acute, pilose; flowers 
ca. 10—15 cm in diameter, 12— 6-merous; calyx 7 3 — J / 2 the length of 
corolla, divided almosttoits middle into acute, lanceolate, pilose lobes; 
petals violet or lilac -purple, stellately spreading, linear -lanceolate, subulate - 
pointed, shortly glandular -pilose, stamens 24— 32, the pubescent filaments 
toward base, the anthers reddish; hypogynous scales straight, flat, 
subquadrate; fruit of 12— 16 divergent green glandular -pilose follicles. 
Fl. July — August, fr. August — September. (Plate II, Figure 6a— d). 

Sandy, calcareous soils, schistose rocks, in the alpine and subalpine 
zones, at 1,300— 2,600 m. Caucasus: Cisc. (often), W. Transc. (rare in 
Abkhazia), E. Transc, Dag. (Samur River). Endemic. Described from the 
Caucasus (Tushetiya). Type in Leningrad. 

Note. S. caucasicum Rupr. is one of the numerous geographic races 
ofS. tectorum L. It differs from S. t e ct o r um in its smaller size 
(45—60 (100) cm in S. t e c t o r um), its smaller fewer -flowered inflorescence, 
and longer petals. S.tectorum is a native of the mountains of central 
and Southern Europe and occurs in the Soviet Union in gardens or, possibly, 
as an escape from cultivation. S. caucasicum Rupr. is represented by 
a series of forms with puberulent or perfectly glabrous basal leaves. 



lo 




PLATE II. 1-Tillaea vaillantii Willd.: a) flower, b) part of shoot with flower and leaves; 
2-T.aquatica L.: a) flower, b) part of shoot with flower and leaves; 3 - T. a la t a Viv.: a) flower, 
b) part of plant with flower and leaves; 4— Umbilicus opposi tif oli us Ldb.: a) part of corolla, 
b) part of inflorescence in fruit; 5-Sempervivum caucasicum Rupr.: a) flower, b) petal with 
stamen, c) pistil, d) sepal; 6 - S. soboli f erum Sims.: a) flower, b) petal with stamen, c) pistil, 
d) sepal. 



17 



21 



22 



In the herbarium of the Botanical Institute of the Academy of Sciences 
there is a specimen of S. flagelliferum Fisch. from Mt. Besh-Tau, 
described by Fisher; it appears that this specimen should also be 
referred to S. caucasicu m. Ledebour reports S.flagelliforme 
Fisch. ex Link, Enum. hort.berol. II (1882) 20.. also quoted in DC, as 
occurring in Siberia, but there are noSempervivum in Siberia. 
S. vermicular e Guldenst. in Reisen I (1787), 102, ex Ldb., Fl. Ross. II 
(1844 — 1846) 190, recorded for the Caucasus (Terek River) (Guldenst.) is also 
problematic and should presumably be referred to S.caucasicum Rupr. 

Economic importance. The leaves are eaten raw. 

4. S. ruthenicum (Koch) Schnittsp. et Lehm. in Flora (1858) 5.— 
S.globiferum Ldb., Fl. Ross. II (l 844-1 846) 1 89 (p. p.) non L.; Boiss., 
Fl. Or. II, 797 p. p. — S. g lob i f e r um subsp. r ut h e ni c u m Koch, Syn. Fl. 
Germ. ed. 2 (1843) 2 89 (in nota ad S.wulfeni).- S. ruthenicum Koch 
ex Shmal'g., Fl. I (1895) 363; Berger in Engl. u. Pr., Nat. Pflzfm. 18a 

(l 930)423; Praeger, An account of the Sempervivum group (1932) 80.— 
Ic: Praeger, 1. c, f. 23; Borisova in Fl. Yugo-Vost., No. 5, f. 409. 

Perennial, 20 — 35 cm high; stems sulcate, sparsely glandular -puberulent; 
rosettes 4— 6(7)cm in diameter, their leaves oblongly obovate-cuneate, 
broadening in the upper third, short -acuminate, densely hirsute on both 
sides, the margin stiffly long-ciliate; cauline leaves remote, alternate, 
oblong-lanceolate, acuminate, hairy on both sides and ciliate ^margined, 
inflorescence loose, corymbiform, 4—10 cm long, ca. 8—10 cm in diameter, 
with long, many-flowered, pubescent scorpioid branches; pedicels 1— 5 mm; 
bracts linear, acuminate, puberulous; calyx green, basally connate, 
ca. 3—4 mm long, pubescent on the outside, with 10—14 oblong -ovate, acute 
lobes, V 4 — V 3 the length of corolla; petals 10 — 14 yellow, stellately spreading, 
distinct, linear, acuminate, long-glandular -pubescent on the outside; stamens 
20—24, shorter than petals, with enlarged pubescent filaments and yellow 
anthers; follicles divergent, oblong-ovoid, with long straight beak, many- 
seeded, glandular -pubescent; hypogynous scales convex, glanduliform; 
seeds oblong-ovoid, grayish brown, slightly more than 0.5 cm long. Fl. 
July — August, fr. from August. 

Sands and sandy soils; sometimes in pine forests.— European part: 
U. V., V.-Don, U. Dnp., M. Dnp., L. Don, BL, Crim. ? (a single specimen 
collected near Simferopol'). Gen. distr. : Bal. -As. Min. (Balkan Peninsula, 
Rumania). Described from "Russia." Type in Berlin. 

5. S.globiferum L., Sp.pl. (1753) 463, p.p.; Boiss., Fl.Or. II, 797, p. p. - 
S.armenum Boiss. Diagn. Ser. II (l 856) 60. — S. b r a u ni i Ldb., Fl. Ross. II 
(1844 — 1846) 190 p.p., non C. Koch. — S. ibericum Fisch in sched. — 

Exs.: Kotschy, No. 385. 

Perennial, 10 — 15 (2 5) cm high; stems densely glandular -puberulent; 
rosettes 2—5 cm in diameter, their leaves oblong-spatulate, somewhat 
enlarged in upper third, abruptly short -acuminate, finely puberulent on 
upper surface, ciliate on the margin, reddening at the tip; cauline leaves 
remote but overlapping, alternate, oblong -ovate, reddening at the tip, 
dilated toward base, acute, pubescent on both sides, the margin ciliate; 
inflorescence contracted, umbellately corymbiform, 2— 5(6) cm in diameter, 



18 



1 — 3 cm long, branching, with short few-flowered pubescent branches; 
pedicels ca. 1 mm long or flowers subsessile; bracts lanceolate, acuminate, 
puberulent, ciliolate; calyx green, basally connate, 5 mm long, pubescent, 
with 12 — 16 lanceolate acute lobes reddening at the tip, corolla 2—2.5 times 
as long as calyx; petals 12—16, yellow-green or yellow, distinct, lance- 
linear, acuminate, stellately spreading, glandular -pubescent on the outside; 
stamens 24 — 32 , shorter than petals with enlarged violet short -pilose filaments 
andyellow anthers; follicles as many as petals, oblique-ovoid, glandular-pilose, 
erect, thebeak recurved, hypogynous scales erect, subquadrate,lamelliform; 
seeds grayishbrown,ca.0.5 mmlong, ovoid. Fl. July— August, fr. from August. 

Rocks and stony slopes, at 1 ,100— 3,000 m, among subalpine and alpine 
plant communities and xerophytic vegetation of southern slopes. Caucasus: 
Cisc. (Mt. Besh-Tau), W. Transc. (Adzhar -Imeretian Range), E. and S. Transc. 
(often). Gen. distr. : Arm. -Kurd. (Kars, Olty). Described from "Russia." 
Type in London. , 

Note. S.globiferum is reported by Linnaeus as collected in 
"in Rutheno" by S. Gmelin. Since S. Gmelin collected only in the Caucasus 
(the other Gmelin, Johann Georg, collected plants only in Siberia where 
Sempervivum does not occur) and as Linnaeus may have believed 
that the Caucasus could also be referred to as Russia, it is conceivable 
that S. globiferum p.p. (S.armenum Boiss.) was described from the 
Caucasus. However, Linnaeus also mentions other characters of European 
species which are not valid for the Caucasian species. As far as the 
Caucasus is concerned, S. g 1 ob i fe ru m is to be interpreted sensu stricto. 

*6. S. glabrifolium A. Bor. sp. nova in Addenda VIII, p. 357. 

Perennial, 10 — 15 cm high; stems glandular -puberulent; rosettes 1—3 cm 
in diameter; their leaves obovate, acute, 1 cm long, 0.5 cm broad, glabrous 
on both sides, with long-ciliate margin; cauline leaves more elongated, 
lanceolate, acute, ca. glabrous on 0.4 cm broad in upper part, glabrous on 
both sides, only the uppermost ones sparsely glandular -pubescent, the 
margin long-ciliate, inflorescence a corymbiform panicle, 3—5 cm long, 
3—4 cm in diameter, with many-flowered scorpioid branches; bracts 
lanceolate, glandular -puberulent; flowers 6— 7 mm long, 10 — 12 -merous; 
pedicels 1—2 mm long; calyx 3 mm long, connate to 7 3 of its length; lobes 
10 — 12, ovate, subobtuse, glandular -puberulent, Y 4 — V 3 the length of corolla; 
petals linear to lance -linear, acute, yellow, 6—7 mm long, infundibularly 
divergent, glandular -puberulent on the outside, entire, with ciliate margin; 
stamens 20— 24. the epipetalous nearly equaling petals, the others shorter 
by anther length; filaments yellow, dilated toward base, with a few hairs 
in lower part; anthers yellow, rounded-oval; hypogynous scales but 0.25 mm 
long, ca. 0.5 mm broad, convex, rounded at the apex; follicles shorter than 
petals, oblique -ovoid, with long, erect, glabrous style as long as the 
ovary, sparsely glandular -puberulent ventrally, glabrous dorsally; follicles 
ovoid-lanceolate, many-seeded, with suberect, slightly recurved beak; seeds 
slightly more than 0.5 mm long, lanceolate, brown. Fl. June, fr. from June. 

Rocks in the districts of Turkey, possibly in USSR territory. 

Gen. distr. : Arm. -Kurd, [former] Artvin District, Chorokh [Coruh] and 
Tsriya rivers). Described from the Artvin District. Type in Leningrad. 



19 



Section 2. JOVISBARBA DC, PI. rar. jard. Gen., No. 21 , obs. (1829) 80; 
Prodr. Ill, 413 (j o v i b a r b a) p. p. ; Koch, Syn. Fl. Germ, (l 837) 264. - 
Jovibarba Opiz, Seznam (1852) 54, pro gen. — Flowers (5)6 -merous, 
campanulate; petals erect, fimbriate. 

7. S. soboliferum Sims in Bot. Mag. (l 812 ) 1457; Shmal'g., Fl. I, 363; 
Berger in Engl. u. Pr., Nat. Pflzfm. 18a (1930) 424; Praeger, An account 
Semperv. group (1932) 100.- S. glob if e rum Rchb. Iconogr. (1834) 
tab. 839, non L. - S. hirtum Ldb.. Fl. Ross. II (1844-1846) 190, non L. - 
Ic: Hegi, III, Fl. IV, 2 (1925) f. 931 h-k; Rchb., Ic. Fl. Germ. XXIII (l 898-1899) 
66; Syreishch., Fl. Mosk. gub. II (1907)216.- Exs.: Fl. exs. austro-hung., 
No. 1307; Meinsh.. Herb. Fl. Ingr. (1866), No. 247. 

Perennial, 10— 25 (40) cm high; stems glandular -pubescent; rosettes 
2—4 cm in diameter, their leaves oblong-cuneate or nearly obovate, 
acuminate, fleshy, bright green, glabrous on both sides, the margin beset 
with long white cilia; cauline leaves oblong or lanceolate, sessile, glabrous, 
acuminate, turning red at the apex, the margin ciliate; inflorescence a 
corymb iform many -flowered dichasium, 5— 7 cm in diameter; branches of 
9 . inflorescence scorpioid; bracts lanceolate; flowers campanulate, 

6-merous; calyx 7— 10 mm long, glandular-pubescent; lobes oval-lanceolate, 
ciliate-margined, reddish-tipped; petals linear -oblong, 12 — 14 (l7)mm 
long, twice length of calyx, pale yellow or greenish, erect, glandular -pubescent 
on both sides, campanulately connivent at apex; stamens 12 shorter than 
corolla; filaments green, sparsely glandular -pubescent; anthers yellow; 
hypogynous scales subquadrate, slightly emarginate. ca. 1 mm long, green; 
follicles straight, gradually tapering into a beak half as long as the fruit. 
Fl. July — August, fr. from August. (Plate II, Figure 6a— d). 

Dry sandy sites, mainly pine forests; also limestones and rocky river - 
banks; sometimes naturalized.— European part: Lad.-Ilm.. U. V., V.-Kama, 
U. Dnp. Gen. distr. : Centr. Eur. (Baltic republics, Poland, Germany, 
Czechoslovakia, Hungary). Described from Europe. Type in London. 

Economic importance. The leaf rosettes are sometimes used as a 
vegetable (Meinshausen). 



Subfamily 4. SEDOIDEAE Berger in Engl, und Prantl, Nat. Pflzfm. 18a 
(1930) 436.— Stamens in two series, as many or twice as many as petals; 
flowers 4—6 (9)-merous; petals distinct or basally connate; leaves of 
variously shaped, alternate, opposite, whorled or rosulate; underground 
organs variable. 



Genus 6 98. RHODIOLA * L. 
L.,Gen.pl.ed.5(l754) 457. 

Flowers 4 - or 5 -merous (exceptionally 6 -merous), usually dioecious, 
rarely bisexual; calyx persistent; corolla yellow, yellowish green, 
cream-colored, whitish pink, or red; inflorescence terminal, corymbiform, 



From the Greek rhod ia rh i za, pink root, or else diminutive of the Greek rhodia orrhodon, rose, 
referring to the odor which is reminiscent of roses. 



20 



capitate -corymbiform or racemose (subgenus Clementsia); follicles 
straight, mostly short -beaked or beakless; seeds numerous, small. 
Perennials with woody, robust, usually branching, multicipital caudex 
covered with congested appressed leaves, these often obsolescent, triangular 
or suborbicular, scalelike, membranous, brown or fuscous; stems few 
to numerous, unbranched, erect or somewhat flexuous, often remaining 
from the previous year; leaves alternate, flat to subterete. A high-mountain 
genus; several species occur in N.Am. Type species R. rosea L. 

1. Inflorescence a raceme; flowers bisexual, white or pink, large, 

ca. 1 cm long (Section Clementsia (Rose) A. Bor.) 

1. R. semenovii (Rgl. et Herd.) A. Bor. 

+ Inflorescence corymbiform or compactly corymbiform-capitate; 
flowers usually dioecious, rarely bisexual, whitish, yellow, cream- 
colored, red or pinkish, sometimes only calyx red u . 2. 

2. Caudex thick, densely surrounded by tufts of persistent old stems; 
annotinous stems few, slender, 1—2 mm in diameter, (3) 10— 15 (20)cm 
high, follicles connate at base. (Section Chamae-rhodiola Schrenk) 
3. 

+ Caudex without stem remnants or, if same old stems occasionally 
present, then not tufted; annotinous stems few, (7)l0— 60 cm long, 
(2) 3— 6 mm in diameter; follicles not connate. (Section Eu-Rhodiola 
Schrenk) 9. 

3. Leaves linear, terete, or linear -lanceolate to oblong, entire; stamens 
usually not longer or sometimes but slightly longer than petiols .... 4. 

+ Leaves oval or oblong-elliptic, usually toothed; stamens exceeding 

petals ' '« 

4. Flowers bright yellow and 4-merous or whitish to pale yellow (whitish) 
and 5-merous; scalelike crown leaves acutely triangular, 3—5 (6) mm 
broad 5. 

+ Flowers red, 5-merous; scalelike crown leaves large, rounded- 
triangular, 6-8 mm broad 18. R.coccinea (Royle) A. Bor. 

5. Flowers 5 -merous, pale yellow; inflorescences many -flowered, rarely 
loose; 1—1.5 cm in diameter (stems 10— 20 (30 ) cm long, 2 mm in diameter 
17. R. pamiroalaica A. Bor. 

+ Flowers usually 4-merous, bright yellow, in few -flowered inflorescence 
0.5—1 cm in diameter; stems 3—10 (15) cm long, 0.5— mm in 
diameter 6. 

6. Stems more or less straight, old stems and caudex often turning black 
or grayish brown; leaves acute, linear; follicles ca. 5 mm long, 
lanceolate. (Urals, Siberia) .... 15. R. quadrifida (Pall.) Fisch. et Mey. 

+ Stems recurved, divergent, old stems and caudex grayish, not turning 
black; leaves subobtuse, oblong or linear -lanceolate, subhorizontally 
recurved; follicles ca. 3 mm long, ovoid .... 16. R. kaschgarica A. Bor. 

7. Scalelike crown leaves large, more than 1 cm long, 1 — 1.5 cm broad. 
Plants (8)12 — 17 cm high; leaves coarsely dentate; hypogynous scales 
quadrate 8. 

+ Scalelike crown leaves not more than 0.4—0.5 cm long, 0.5—0.7 cm 
broad. Plants 3—5 (l0)cm high; leaves denticulate or entire; calyx 
yellow; hypogynous scales elongated 19. R. gelida Schrenk. 



21 



8. Plants (8) 12 — 15 cm high; leaves ovate, 0.8—1 cm long, 0.2—0.3 cm 
broad, dark green; calyx red; scalelike leaves 1 cm long and 1 cm 
broad; old stems numerous; follicles with beak less than 1 mm 

long 20. R. recticaulis A. Eor. 

+ Plants 10— 17 cm high; leaves oblong-elliptic to lanceolate, 1 — 1.5 cm 
long, 0.3—0.5 cm broad, light green; calyx yellow; scalelike leaves 
larger, 1 — 1.5 cm long and as broad; old stems few; follicles with 
beak 1 — 1.5 mm long 21. R. litvinovii A. Bor. 

9. Flowers bisexual, sometimes dioecious, mostly 5-merous, less 
often 4-merous), white, pinkish or yellow, 4— 8 mm long; leaves 

linear or lanceolate to oblong 10. 

+ Flowers dioecious, mostly 4-merous (less often 5 -merous), yellow or 
red, 3—6 mm long; leaf shapes various 11. 

10. Flowers yellow, 4—5 mm long, 4 (5)-merous; follicles green, ca. 7 mm 
long; leaves 1—2 mm broad, 1—2 cm long, linear, entire or with 1 or 

2 teeth 15. R. komarovii A. Bor. 

+ Flowers whitish or pinkish, 7— 8 mm long, 5-merous; follicles 

7—10 mm long, red at maturity; leaves 1.5— 3 mm broad, 0.8—2 cm 

long, linear to lanceolate -oblong, entire (Altai) 

13. R. algida (Ldb.) Fisch. et Mey. 

11. Leaves linear to lance -linear, entire or remotely dentate or coarsely 
pinnate-dentate; stems few, tall, 25— 60 cm long 12. 

+ Leaves rounded -cordate to lanceolate, dentate at apex or serrulate, 
rarely subentire; stems mostly in groups from a single shoot 
(7) 10-30 (5)cm long 15. 

12. Leaves alternate or in pseudo-whorls of 3, coarsely pinnate -dentate, 
cuneate; rhizome 0.5 — 1.5 cm long; inflorescence dense, corymbiform 
or umbellate, surrounded by leaves, 1.5—3 cm in diameter; flowers 
yellow or cream -colored 13. 

+ Leaves alternate, subentire, linear -lanceolate, basally broadened; 
caudex robust, elongated; flowers brick-red or if yellow then 
inflorescence spreading, 7—10 cm in diameter, and pedicels 
elongated 14. 

13. Flowers cream-colored, 5— 6 mm long; petals linear -lanceolate; 
stamens as long as or slightly longer than corolla; hypogynous 
scales subquadrate 12. R. stephani (Cham.) Traut. et Mey. 

+ Flowers yellow, 6 mm long; petals lanceolate or oblong -lanceolate; 
97 stamens 1.5 times as long as corolla; hypogynous scales elongated, 

longer than broad (Dau., Ang. -Say.) 11. R. pinnatifida A. Bor. 

14. Flowers yellow; pedicels elongated; inflorescence spreading, 
oligophyllous; leaves lance-linear to linear, 4— 6 (7) cm long, 
0.2—0.5 cm broad, subentire or remotely denticulate; calyx half the 
length of corolla; stamens about as long as or slightly longer than 

petals, with yellow filaments and anthers . 

9. R. kirilowii (Rgl.) Maxim. 

+ Flowers brick-red; pedicels shorter than flowers; inflorescence 

compact, leafy; leaves linear -lanceolate, subentire or very remotely 
dentate at the apex, basally broadened, 2—5 cm long, 0.3—0.7 cm 
broad; calyx 2 / 3 the length of corolla; stamens 1.5 times as long as 

porolla, with red filaments and yellow anthers 

10. R. linearifolia A. Bor. 



22 



28 



15. Stem leafless in upper part; inflorescence dense, corymbiform- 
capitate; stamens twice as long as corolla, with red or green 
filaments; leaves triangular -ovate, amplexicaul, scattered along 
the stem, sometimes the upper more elongated, glaucous; flowers 

red R. heterodonta (Hook, et Thorns.) A. Bor.* 

+ Stem leafless near inflorescence; inflorescence usually surrounded 
by leaves; stamens as long as or slightly longer than corolla, the 
filaments yellow or if red then flowers also red 16. 

16. Flowers red, anthers red 17. 

+ Flowers yellow, anthers green 18. 

17. Filaments red; leaves scattered along the stem, usually glaucous, 
cuneate, oblong-elliptic; flowers 3— 6 mm long. (Far East, E. Arc. : 
An., Chuk.) 8.R. atropurpurea (Turcz.) Trautv. et Mey. 

+ Filaments yellow; leaves imbricately approximated, green, rounded - 
cordate at base of the stem, more elongated above; flowers 2.5—3 mm 
long 6. R. borealis A. Bor. 

18. Leaves oblong -spatulate, elongate -rhomboid, cuneate, sometimes 
subpetiolate, remotely and coarsely dentate at the apex, entire 

below (Sakhalin) 5. R. sachalinensis A. Bor. 

+ Leaves rounded-ovate to elliptic or lanceolate, with broad base, 

crenate, shallowly dentate-serrate or entire, rarely irregularly and 
coarsely dentate 19. 

19. Leaves orbicular to oblong-ovate, imbricately approximated or 
dense; flowers longer than pedicels 20. 

+ Leaves elliptic to lanceolate, remote, entire or remotely denticulate - 

serrulate; flowers shorter than pedicels 2. R. rosea L. 

20. Plants 15—20 (30)cm high, 4— 5 cm in diameter; scalelike crown, 
leaves 5—7 mm long, 5 mm broad, green, oblong -ovate, with broad 
cuneate base, coarsely and irregularly dentate, dense but not 
imbricately approximated 4. R. iremelica A. Bor. 

+ Plant 7— 10 (15) cm high, stem 2 — 3 mm in diameter; scalelike crown 
leaves 4 mm long, 3 mm broad; leaves rounded -ovate, glaucous, 
amplexicaul, entire or remotely dentate; imbricately approximated. 

(Novaya Zemlya, Vaigach Island, Kola Peninsula) 

3. R. arctica A. Bor. 



Section 1. CLEMENTSIA (Rose) A. Bor. - C 1 e m e n t s i a Rose, Bull. 
N. Y. Bot. Gard. 3 (1903) 3, pro gen. — Inflorescence a dense spieiform 
raceme; flowers bisexual, 5-merous, white or pink, ca. 1 cm long. 

1. R. semenovii (Rgl. et Herd.) A. Bor. comb. nova. — S edum semenovii 
Masters, Gard. Chron. X (1878) 267; Berger in Engl. u. Pr., Nat. Pflzfm. 18a 
(1930) 443. — Umb ili cus semenowii Rgl. et Herd, in Bull. Soc. Nat. 
Mosc. XXXIX (1866) 65.- U. linif olius Ost.-Sack. et Rupr. in Mem. Ac. 
Sc. Petersb., Ser. 7, XIV (186 9) 46. - U. 1 i n e a r if o 1 iu s A. Franchet in 
Ann. Sc. Nat. Ser. VI, XVI (1883) 290.- Cotyledon semenovii O. et 
B. Fedtsch., Consp. Fl. Turk. 3 (1909) 70.- Ic. : Praeger in Journ. Royal 
Hort. Soc. (1920-1921) f. 27; Frod. in Acta Horti Gothoburg. (l 930) 
textfigs 134-140? 

* R.viridula A. Bor. (in Addenda VIII, p. 396 should be distinguished: light green, with yellow, usually 
5-merous flowers, light green leaves, larger fruit, etc.- T. Sh.(Chatkal, Talass Ala-Tau,and Bol'shoi 
Chimgan ranges). Described from the Chatkal Range. Type in Leningrad. 

23 



29 



Perennial; caudex thick, branching, upper part covered with scaly 
triangular leaves ca. 5 mm long, 4 mm broad, gradually passing into green 
cauline leaves; stems few, 35— 60 cm high, 0.5— 0.6 cm in diameter, rounded, 
densely leafy, erect, simple; leaves linear, entire or remotely dentate, 
acute, the lower leaves to 1 cm long and 2 mm broad, the middle leaves to 
7 cm long and 3 mm broad, the upper to 3.5 cm long and 2 mm broad; 
inflorescence a long, dense, spikelike raceme; flowers 5 -merous, bisexual, 
short -pediceled or sessile, ca. 1 cm long, with long-linear bracts; sepals 
green, linear, acute, 2 / 3 as long as petals; petals white or pink, lanceolate; 
stamens 10, straight, as long as petals, with white filaments and red 
anthers; hypogynous scales quadrate, small; follicles straight, 1 cm long, 
with long slender beak greenish white, turning red, thus reddening the 
inflorescence; seeds 1 mm long, ovoid, alate. June — July. 

Moist, stony soils, riverbanks, mountain forests, passes, mossy alpine 
meadows and bogs up to 3,500 cm. — Centr. Asia: T. Sh., Pam.-Al. Endemic. 
Described from the Trans -Hi Ala-Tau. Type in Leningrad. 



Section 2. EU-RHODIOLA Schrenk in Fisch. et Mey., Enum.pl. nov. 
(1841 ) 6 7.— Caudex without remnants of stems or with but one old stem, 
not tufted; stems few, (2)3—6 mm in diameter; follicles apocarpous. 



Series 1. Roseae Praeger, 1. c, 28, ex parte, pro grege. — Flowers yellow, 
greenish, or red, dioecious, small, 3— 4 mm long, 4 -merous; leaves rounded - 
ovate to lanceolate. 

2. R. rosea L., Sp.pl. (1753) 1035; Small, North. Amer. FL, v. 22, p. 1 
(1905) 57.- R. elongata Fisch. et Mey. in Schrenk, Enum. pi. nov. I (l84l) 
83; Middend., Fl. Ochot. (1856) 39. - Sedum roseum Scop., Fl. Cam., 
ed.2,1 (1772)326; Kryl., Fl. Zap. Sib. VI, 1407; Berger in Engl. u. Pr., Nat. 
Pflzfm. 18a (1930). 440.- S.rhodiola DC, PL Grass. (1805) 143; Prodr. 
111,401.- S. a It ai cum G., Don, Gen. Syst. gard. and Bot. Ill (1834) 114.- 
S. elongatum Ldb., Fl. Ross. II (1844-1846) 178 (non Wallich); Turcz., 
Fl. baic.-dah. I, 434.- Ic: DC, 1. c, tab. 143, 144; Rchb., Ic. Fl. Germ. 
XXIII (1898-1899) tab. 41; Praeger in Journ. Hort. Soc. XLVI, f. 5b; Frod., 
Acta Horti Gothoburg. V (1930) f. 102-107. 

Perennial; root thick, straight; caudex short, thick, with scalelike, acute, 
triangular, membranous leaves, 5 mm long, 4 mm broad; stems few, 
25—30 cm, on moist soils sometimes to 50 cm high, 4—6 mm in diameter; 
leaves entire, at tip declinately serrate, with few teeth, acute or acuminate, 
0.7—3.5 cm long, 0.5 — 1.5 cm broad; inflorescence dense, corymbiform, 
many -flowered, 3—4 (6) cm in diameter, 2 cm long, with elongated, forked, 
curved slender branches; flowers shorter than pedicels, dioecious, 4- rarely 
5 -merous, yellow, 3—4 mm long; sepals lance-linear, ca. 1 mm long, 
V 4 — Y 2 length of petals, obtuse, yellow or greenish; petals linear or oblong, 
subobtuse, yellow or greenish; stamens exceeding the petals; filaments 
and anthers yellow; hypogynous scales 2—3 times as long as broad, 
1 — 1.5 mm long, 0.5 mm broad, slightly attenuate, truncate, emarginate; 



24 



follicles curved, greenish, 6-8 mm long, erect, lanceolate or linear- 
lanceolate, with short, slender beak ca. 1 mm long; seeds 2 mm long, 
lanceolate. May — June. 

Arcto -montane plants of moist soils, riverbanks, pebbles, cliffs and 
pine forests, rock crevices, rocky slopes, dunes, sandy soils in tundra.— 
Arctic: Arc. Eur., Chuk., An.; European part : Dv. -Pech., V. -Kama.; 
W.Siberia: Alt.; E.Siberia: Yenis., Dau., Lena-Kol.; Centr.Asia: 
Dzu. -Tarb. Gen. distr.: mountains of W. Eur., Bal.-As. Min., Mong., 
China (Shansi), Dzu.-Kash., Scand. Described from Europe. Type in 

Leningrad. 

Note. An extremely polymorphic, widely distributed species. A 
number of authors (Regel and Tiling, Maximowicz) have recognized a 
series of distinct forms. We separate from R. r o s e a s.lat.: 
R.arct ica, R. ir emelica, and R. s a c ha 1 i n e n s i s as species which 
have undoubtedly evolved from the widely distributed arcto -montane 
R. ro s ea, under particular ecological conditions. R. bo real is A. Bor. 
is intermediate between R. r os ea s. lat. and R. a t r o p u r pu r e a S.lat. 

Economic importance. Used medicinally; pharmaceutical name: 
Radix Rhodia. 

3. R. arctica A. Bor. sp. nova in Addenda VIII, p. 357.- R. sibir ica 
Sweet ex Ldb., Fl. Ross. II (1844-1846) 179, p. p.- R. rosea Fisch. et 
Mey. ex Schrenk, Enum.pl. nov. I (l84l) 67 (non L.).- Sedum rhodiola 
Ldb., I.e., 179.- S. rhodiola var. 1 at i f o 1 i a Rgl. in Trautv., Fl. Nov. 
Semi. (1871) 24.- Ic. : Praeger in Journ. Royal Hort. Soc. (1921 ) f.4; 
Frod. in Acta Horti Gothoburg. V (1930) f. 108-113. 

Perennial; roots thick, cordlike; caudex short, 4-5 cm long, sometimes 
branching, thick, with scalelike, acuminate, rounded -ovate leaves 4 mm long, 
3 mm broad; stems 7-10 cm high, 2-3 mm in diameter, slightly curved, 
often numerous; leaves glaucous -green, imbricately approximated, rounded - 
ovate, acuminate, entire or remotely serrate at apex, amplexicaul; 
inflorescence many -flowered, compact, ca. 2 cm in diameter, 1 cm long, 
leafy; flowers 4-merous, dioecious, 3 mm long, longer than pedicels; 
sepals lanceolate, V4-V2 the length of petals, subobtuse, yellow or green; 
petals lanceolate, carinate, obtuse, yellow, 3 mm long; stamens 8, exceeding 
the petals, with bright yellow filaments and rounded anthers; hypogynous 
scales ca. 1 mm long, quadrate or twice as long as broad, emarginate; 
ovaries straight, obsolete in staminate flowers; follicles ovoid, green, 
with short beak, 4-5 mm long; seeds less than 1 mm long, oblong. Fl. 
July -August, fr. August - September. (Plate III, Figure la-b). 

Stony and lichen tundra, shallows, rock crevices, clayey soils. - 
Arctic: Nov. Z, Vaigach Island, Kola Peninsula. Endemic. Described 
from Novaya Zemlya. Type in Leningrad. 

Note. Closely related to R. ro s ea, from which it is distinguished 
by its height, the glaucous -green coloring, shape and arrangement of 
leaves, length of flowers and pedicels, and shape of hypogynous scales. 

4. R. iremelica A. Bor. sp. nova in Addenda. VIII, p. 358.- Sedum 
roseum auct. p. p. — S. r h o d io 1 a auct. p. p. 

Perennial; root thick, straight; caudex branching, robust, covered in its 
upper part with scalelike, membranous, triangular, acute leaves, 5— 7 mm 



25 



32 



long, 5 mm broad (the upper more elongated); stems usually few, 15— 20(30)cm 
high, 4—5 mm in diameter; leaves approximate, green, oblong-ovate, 2—2.5 cm 
long, 1—1.5 cm broad, acute, broadly cuneate -based, the whole margin 
coarsely and irregularly dentate; inflorescence dense, corymbiform, 2—4 cm 
in diameter, 1 — 1.5 cm long, its branches straight, thickish, not forked, 
few -flowered in pistillate plants, slender and branching in staminate plants; 
flowers 4 -merous, dioecious; sepals of pistillate flowers linear-lanceolate, 
acute, ca. 1 mm long, in staminate flowers linear, subobtuse, ca. 1.5 mm long; 
petals green, linear, subobtuse, ca. 3 mm long; stamens slightly longer than 
petals, with rounded anthers and filiform filaments; follicles 5— 7 mm long, 
oblong-lanceolate, thickish with short, divergent beak, ca. 0.5 mm long 
(or shorter); seeds lanceolate, slightly longer than 1 mm. Fl. June, fr. July.- 

On summits of bald mountains, rocky outcrops, shady sites. — European 
part: V. -Kama, S.Urals (iremel', collected by Shell, Litvinov, Tyulina), 
Bashkir ASSR: Abzelilovskii District, Krykty-Tau Range at the latitude 
of Lake Bannoe (collected by Krasheninnikov). Endemic. Described from 
Irmel'. Type in Leningrad. 

5. R. sachalinensis A. Bor. sp. nova in Addenda VIII, p. 360.— Sedum 
r o s e u m auct. p. p.— S.rhodiola auct. p. p., non DC. 

Perennial; root robust, cordlike, straight; caudex 3—6 cm long, 1—3 cm 
broad, densely covered with scalelike imbricate, oblong -triangular, acuminate 
leaves, 0.6 cm long, 0.3 cm broad, gradually passing into green leaves; 
stems 2—5, 10—15 (20)cm high, 3—4 mm in diameter; leaves alternate, 
oblong -spatulate, elongate -rhomboid, 2—2.5 cm long, 0.5—0.7 cm broad, 
cuneate, abruptly narrowing, coarsely dentate in upper part, entire in 
lower part, acute; inflorescence many -flowered, loose, 2—2.5 cm in diameter, 
surrounded by leaves; flowers yellow, small, dioecious, 4 -merous, rarely 
5 -merous; sepals 2 / 3 the length of petals, acute, oblong; petals oblong- 
lanceolate, subobtuse, 3 mm long; stamens slightly longer than petals, with 
yellow filaments and rounded anthers; hypogynous scales emarginate, 
slightly longer than broad; follicles 4—5 mm long, lanceolate, with short 
beak; seeds ovoid, ca. 1 mm long, 0.5 mm broad. June — August. (Plate III, 
Figure 2a— b). 

In rock crevices and stony sites.— Far East: Sakh. Gen. distr. : Jap.-Ch. 
(Kurile Islands). Described from Sakhalin. Type in Leningrad. 

6 . R. heterodonta (Hook, et Thorns. ) A. Bor. comb. nova. — Sedum 
heterodontum Hook, et Thorns, in Journ. Lin. Soc. Bot. II (1858) 95; 
Berger in Engl. u. Pr., Nat. Pflzfm. 18a (1930) 430. - S. c r enulatum 
Hook, et Thorns. (? ) in Journ. Linn. Soc. Bot. II (1858) 96. — Ic. : Praeger in 
Journ. Royal Hort. Soc. XLVI (l 920-1 921 ), f. 7; Frod. in Acta Horti Gothoburg. 
(1930) pi. XIII, 2 (sub var.). 

Perennial; root robust, vertical, with insignificant branching; caudex 
branching, covered with scalelike subobtuse -triangular leaves 7— 8 mm 
long, 8 mm broad; stems erect, 1—3, usually 30—40 cm, less often up to 10 cm 
high, 4— 5 mm in diameter; leaves remote, triangular -ovate, broadly 
cordate, sometimes the upper elongate, sessile, amplexicaul, coarsely 
dentate, usually glaucous; inflorescence compact, capitate -corymbiform, 
not surrounded by leaves, 1.5—2 cm in diameter, 1—1.5 cm long; pedicels 



26 



short; flowers small, dioecious, 4 -merous, 3-4 mm long (without stamens); 
sepals reddish or greenish, 1.5 times as long as calyx; stamens twice as 
long as petals or longer, their filaments reddish or greenish, ca. 5 mm 
long, the anthers yellow or reddish; the 4 episepalous stamens somewhat 
shorter than the other 4; pistils with short, thickish styles, lanceolate; 
hypogynous scales oblong or subquadrate, emarginate, bright red or 
orange, half the length of fruit; follicles as long as petals, straight, 
linear-oblong, with short, reflexed beak; seeds brown, elliptic, 1.5 mm 
long. May- June. (Plate III, Figure 3a). 

Stony soils, in mountains, up to 4,000 m. - Centr.Asia: T. Sh., Pam. -Al. 
Gen. distr. : Ind.-Him. (NW India and Him.), Mong. (Centr. Mong. and Tib.), 
Dzu.-Kash., Iran. (Afghanistan). Described from the Himalayas. Type in 
London. 

7. R. borealis A. Bor. sp. nova in Addenda VIII, p. 360. - Sedum 
roseum auct. p. p. - S. r h o d i o 1 a auct. p. p. 

Perennial; roots cordlike, long, little branching; caudex branching, 
1-4 cm broad and as long, its upper part covered with subobtuse, triangular- 
ovate, imbricated, brown scalelike leaves 4-5 mm long, 3-5 mm broad; 
stems 5— 10 (15) cm long, 2-3 mm in diameter, numerous, densely leafy, 
slightly reflexed before anthesis; leaves rounded -cordate, entire, acute 
and subopposite in lower part of stem, upper leaves alternate, more 
elongated to oblong, subacute, remotely dentate near the apex; inflorescence 
dense, compact, rounded, many -flowered, 1.5-2 cm in diameter, 1-1.5 cm 
long; flowers dioecious, 4 (5)-merous, small, 2.5-3 mm long; sepals acute, 
lanceolate, half the length of petals, red, sometimes yellow at base; petals 
2.5-3 mm long, carinate, red or yellowing; stamens 8-10, barely longer 
than petals, with yellow filaments and rounded, yellow anthers; stamens 
sometimes 4-5, in 1 whorl; pistils with erect style and clavate stigma 
obsolete in staminate flowers; hypogynous scales half as broad as long, 
tapering toward base, with deep apical notch; follicles 4 mm long, ovoid, 
with short, clavate beak; seeds 1-1.5 mm long, 0.5 mm broad, ovoid, grayish 
brown. June— August. 

Tundra, seashores, steep stony and dry slopes, on coarse clayey skeletal 
soils. - Arctic: Arc. Sib. (Yenis., Yakutsk area, Dikson Island). Endemic. 
Described from near the village of Dudinka. Type in Leningrad. 

Note. This species is probably of ancient hybrid origin, a cross 
between R. rosea and R.atropurpurea but with fixed hereditary 
properties and a definite distribution area. 

8. R. atropurpurea (Turcz.) Trautv. et Mey. in Middend., Fl. Ochot. (1856) 
39.- Sedum a t r o pu r p u r eum Turcz. in Bull. Soc. Mosc. I (1840) 1 3, 70; 
Ldb., Fl. Ross. II, 179; Kom., Fl. Kamch. II, 200. - Sedum roseum var. 
atropurpureum Praeger in Berger in Engl. u. Pr., Nat. Pflzfm. 18a 
(1930) 440. - Ic. : Praeger in Journ. Royal Hort. Soc. (1921 ) f. 5c. 

Perennial; root vertical, few -branched; caudex elongated, covered with 
scalelike oblong, acute leaves 5-6 mm long, 3-4 mm broad; stems numerous, 
somewhat reflexed, 10-20 (25) cm high, 2-5 (7) cm [sic] in diameter; leaves 
scattered along the stem, cuneate, usually acute, denticulate, coarsely 
dentate or subentire, 1 .5-3 cm long, 0.5-0.8 cm broad, oblong-elliptic, 



27 



usually glaucous; inflorescence ca. 2—5 cm in diameter, ca. 1 .5— 2.5 cm 
long, surrounded by leaves; flowers dark purple, shorter than or as long as 
q4 pedicels, 3—6 cm long, dioecious, sometimes bisexual, 4- or 5 -merous; 
sepals red, linear or lanceolate, acute, broadening toward base, 1.5—2 or 
4—5 mm long; petals oblong-elliptic or lanceolate, obtuse, 3—4 or 5—6 mm 
long; stamens with red filaments and rounded yellow or yellow -red anthers, 
as long as or slightly longer than petals; hypogynous scales oblong or 
quadrate, emarginate or nonemarginate, dark red, thickened; follicles 
ca. 8 mm long with long straight or slightly reflexed styles, oblong -ovoid; 
seeds 1—2 mm long, lanceolate or oblong -ovoid, dark brown. May — June. 

Taluses, rocks, gravels, river valleys; singly or in groups.— Arctic: 
An.,Chuk.; Far East: Kamch., Okh., Uda. Gen. distr. : N. Am., Alaska, 
Rocky Mountains (?). Described from the Okhotsk territory. Type in 
Leningrad. 

Note. Highly polymorphic, undoubtedly a collective species, but owing 
to lack of adequate material it cannot be separated into its components. 

Clearly differentiated specimens have been collected in the vicinity 
of the port of Ayan and the Okhotsk coast near Taui Bay; f. ochotensis 
A. Bor. is characterized by larger 5—6 mm long flowers, may-flowered 
inflorescence, quadrate hypogynous scales, shape of leaves which are 
predominantly coarsely dentate, etc. 



Series 2. Linearifoliae A. Bor. Flowers yellow, cream -colored or 
red, 3—6 mm long, 4 - or 5 -merous, dioecious, less often bisexual; leaves 
linear to linear-lanceolate. 

9. R. kirilowii Rgl. ex Maxim., Pr. Fl. Amur. Suppl. Ind. Fl. Pekin. (1859) 
472.- Sedum kirilowii Rgl. in Rgl. et Tiling, Fl. Ajanensis (1858) 92 in 
adnot., No. 114; Maxim. Bull. Acad. Petersb. 29 (1883) 128; Fedtsch. O. et 
B., Consp. Fl. Turk. 3 (1909)72; Berger in Engl. u. Pr., Nat. Pflzfm. 18a 
(1930) 440.- S. elongatum Kar. et Kir., Enum.pl. Song. (1841 ), No. 360; 
Ldb. Fl. Ross. II, 178 p. p.- Ic. : Praeger in Journ. Royal Hort. Soc. (1921 ) f. 8; 
Frod. in Acta Horti Gothoburg. (1930) pi. XIV, textfig. 133. 

Perennial; root thick, vertical; caudex thickened, 1.5 cm broad, in its 
upper part, with scalelike 4 mm long, 2 mm broad triangular, acute membranous 
leaves; stems few (l— 2), (15)40—50 cm high, 4—5 mm thick, densely leafy; 
leaves alternate, sessile, linear to linear-lanceolate, 4—6 (7) cm long, 
0.2—0.5 cm broad, remotely dentate, rarely subentire, acute; inflorescence 
corymbiform, many -flowered, spreading, 7—10 cm in diameter; flowers 
dioecious, 4- or 5 -merous, 3 mm long; staminate flowers shorter, pistillate 
flowers longer than pedicels; sepals linear, acute, half the length of petals; 
...j. petals linear-lanceolate, broadened in upper part, greenish yellow; stamens 
as long as or longer than petals, with yellow filaments and anthers; pistils 
as long as petals; hypogynous scales oblong, twice as long as broad, 
emarginate or subobtuse, yellow, about as long as pistil; follicles 4—5 mm 
long, straight, green, with short beak; seeds 2 mm long, ovoid. March— May. 

Rock crevices, slopes, passes. — Centr.Asia: T. Sh. (Narym Range, 
Terskei Ala-Tau), Pam.-Al. (Alai Range). Gen.distr.: Jap. -Ch. (N. China), 
Tib., Dzu. -Kash. Described from Peking, after specimens collected by 
Kirilov. Type in Leningrad. 

Economic importance. Ornamental; overwinters very well in Leningrad. 



28 



10. R. linearifolia A. Bor. sp. n. in Addenda VIII, p. 3 60.- Sedum 
kirilowii var. linifolium Rgl. et Schmalh. in Acta Horti Petrop. V 
(1877) 583.- S. linifolium rubrum hort.; Rhodiola linifolia 
rubra hort.— Ic: Rgl., Gartenfl. t. 1080 sub S. ki r ilow i i; Praeger in 
Journ. Royal Hort. Soc. (1921 ) f. 9. 

Perennial; root thickened, caudex robust, covered in upper part with 
scalelike, triangular, acuminate leaves, 0.6 mm long, 0.5 mm broad. Dark 
green plants with 1-3 stems 25-30 cm high, 4-6 mm thick, sulcate; leaves 
alternate, linear -lanceolate, broadened toward base, 2-5 cm long, 3-7 cm 
broad, with very sparse teeth confined to the apex, or else entire, sessile, 
acuminate; inflorescence corymbiform, many-flowered, compact, usually 
leafy, 1.5—5 cm in diameter, 1.5 cm long; flowers dioecious, sometimes 
bisexual, mostly 5, rarely 4-merous; pedicels shorter than flowers; sepals 
linear, acuminte, 2 / 5 the length of petals, greenish; leaves linear-lanceolate, 
4 mm long, subobtuse, brick red; stamens 1.5 times as long as petals, with 
red filaments and bright yellow anthers; hypogynous scales subquadrate, 
emarginate, half the length of pistils; follicles 6-8 mm long, 1.5-2 times 
as long as petals, with short beak. May— July. 

Forest meadows, forest edges, on rocks up to 3,000 m. — Centr. Asia: 
T. Sh. (Narym, Aleksandrovskii [Kirghiz], and Trans -Hi Ala-Tau ranges. 
Gen. distr.: Dzu.-Kash. Described from the vicinity of Alma-Ata, 
Bozyngen Pass. Type in Leningrad. 

Note. R. linear if olia A. Bor. has been confused with R. k i r i 1 ow i i 
Rgl., described from the vicinity of Peking, and known to occur in the 
Chinese provinces of Kansu, Szechwan, and Chihli [Hopeh], in Tibet (Kham 
Province), in Kashgaria, and in the USSR in the Narym, Terskei Ala-Tau, 
Karakol, and Alai ranges. In the Soviet Union the two species occur 
together; however, R. 1 i n e a r i fo 1 ia does not penetrate further than 
Kashgaria. Praeger has obtained hybrids of the two species with pale 
orange flowers or with yellow petals, sepals, and fruitlets and with dark red 
filaments. The same species also appear to form hybrids in nature. 

Economic importance. A popular ornamental species. 

11. R. pinnatifida A. Bor. sp. nova in Addenda VIII, p. 361.— Sedum 
d e nt at um Steph. p. p. in herb., non DC. 

Perennial; root cordlike, little branching; caudex short, 1 — 1 .5 cm 
long, 1.5-2 cm broad, covered with scalelike, oblong -triangular, acute 
brown leaves, 4-5 mm long, 3-4 mm broad; stems 2-4, arising from each 
branching of the caudex, 15-20 cm high, 3-4 mm in diameter, erect, densely 
leafy; leaves lanceolate to linear -lanceolate, coarsely, deeply pinnately, 
and irregularly dentate, alternate, in pseudo -whorls of three, 2—3.5 cm 
long, 3—5 mm broad, gradually tapering toward base, acuminate; inflorescence 
dense, umbelliform, many -flowered, surrounded by leaves, 2.5 cm in 
diameter, 1.5 cm long; flowers 4-merous, dioecious, smaller than pedicels; 
sepals 4 mm long, 2 / 3 the length of petals, acuminate, lanceolate, yellow -green; 
petals 6 mm long, lanceolate, or oblong-lanceolate, yellow, subobtuse; 
stamens 8, 1.5 times longer than petals, with yellow filaments and rounded 
yellowish-greenish anthers; hypogynous scales elongated, 1 mm long, 0.5 mm 
broad; follicles ca. 7 mm long, oblong -lanceolate, tapering into a beak; 
seeds oblong, slightly more than 2 mm long, ca. 1 mm broad, subobtuse, 
brown. Fl. June — August, fr. July — August. 



29 



37 



Gravelly, argillaceous, moistened soils, often near riverbeds, in 
floodplains, on alpine meadows in bald mountains. E. Siberia: Dau., 
Ang. -Say. Gen. distr. : Mong. Described from Transbaikalia 
(Mt. Sokhondo). Type in Leningrad. 

Note. See note on R. s t ep ha ni (Cham.) Trautv. et Mey. 

12. R. stephani (Cham.) Trautv. et Mey. in Middend., Fl. Ochotens. (1856) 
39.- Sedum stephani Cham. Linnaea VI (1831 ) 549 p. p.; Ldb., Fl. Ross. 
II, 178; Berger in Engl. u. Pr., Nat. Pflzfm. 18a (1930) 442.- S. dent at urn 
Steph. p. p. ex Cham., 1. c. — C h a m a e r h o d i o 1 a stephani Nak., Rep. first 
Scient. exped. to Manchoukuo IV (1934) 31.— Ic: Praeger in Journ. Hort. 
Soc. (1921) f.23. 

Perennial; root thickened, cordlike, branching; caudex short, few - 
branched, ca. 0.5—1 cm broad, covered with scalelike, triangular, 
subobtuse leaves, 5 mm broad, 5 mm long; stems few, 10—25 cm high, 
ca. (2.5) 5 mm in diameter; leaves lanceolate to linear -lanceolate, 3—5 cm 
long, (3)6— 8 mm broad, acuminate, coarsely and deeply dentate, cuneate at 
base, entire below, pale green; inflorescence dense, (l.5) 2—3 cm in diameter, 
leafy, with few branches; flowers 4 (5)-merous, dioecious, pedicels shorter 
than flowers; sepals linear, Y 2 — 2 / 3 as long as petals, greenish yellow; petals 
cream -colored or whitish, obtuse, linear -lanceolate, 5— 6 mm long; stamens 
as long as or slightly longer than petals, with pale yellow, whitish filaments 
and pale yellow, sometimes pinkish anthers; stamens opposite to petals, 
shorter than petals; ovaries lanceolate, gradually broadening toward base, 
with elongated erect style and thickened stigma; hypogynous scales 
subquadrate; follicles green or reddish, oblong-lanceolate, erect, 7— 10 mm 
long, with straight beak, ca. 1 mm long; seeds obovoid, ca. 2 mm long, brown. 
Fl. June — August, fr. July — August. 

Forest edges, moist forests, seashores, river valleys.— Far East: Okh., 
Uda. Endemic. Described according to specimen from Willdenow's 
herbarium, collected by Redovskii as No. 8910, and bearing the erroneous 
indication of Kamchatka, where Redovskii had never been and where 
S. stephani does not occur. Type in Leningrad. 

Note. Chamisso considered Sedum dentatum Steph. in herb, 
from Transbaikalia as identical with the Okhotsk species. The appellation 
S. dentatum Steph. could not be retained, for there already existed a 
S. dentatum DC. in the Caucasus, therefore Chamisso was undoubtedly 
wrong in describing S. stephani as an Okhotsk and Transbaikalia species. 
R. stephani s. str. is characteristic for the Far East. A closely related 
species growing in Transbaikalia, R.pinnatifida A. Bor., is differentiated 
by color of corolla (yellow flowers, cream-colored or whitish in 
R. stephani), color of leaves, longer stamens, shape of hypogynous scales, 
and a number of other characters. 



Series 3. Algidae A. Bor.— Flowers bisexual, 5 -, less often 4 -merous; 
leaves linear to oblong -lanceolate. 

13. R. algida (Ldb.) Fisch. et Mey. in Schrenk; Enum. pi. nov. I (l84l) 70, 
Sedum algidum Ldb., Fl. alt. II (l 830) 1 94; Fl. Ross. II, 177; Kryl., Fl. 
Zap. Sib. VI, 1406; Berger in Engl. u. Pr., Nat. Pilzfm. 18a (1930) 442.- 



30 



S. euphorbioides Schlecht. ex Ldb., Fl. Ross. II (1844-1846) 177.- 
Chamaerhodiola algida Nakai, Rep. of the first Scient. exped. to 
Manchoukuo, Sect. IV (1934) 30.- Ic: Ldb., Ic. PL Fl. Ross. V (1834) 
tab. 418. 

Perennial; root thick, long; caudex multicipital, covered with scalelike, 
triangular, acute leaves; stems numerous, erect or spreading, densely 
leafy, 6—18 cm high, 1.5—2.5 mm in diameter; leaves alternate, sessile, flat, 
linear -oblong or linear, entire, bluntly acuminate, 8—20 mm long, 1.5—3 mm 
broad; inflorescence a dense simple corymb; flowers 5-, rarely 4-merous, 
usually bisexual, sometimes pistils obsolete, about as long as pedicels; 
sepals ca. 4 mm long, reddish, linear -oblong, obtuse; corolla large, 
1.5—2 times the length of calyx; petals ovate -lanceolate, bluntly acuminate, 
7— 8 mm long, white or dingy pink; stamens 10, slightly shorter than or 
about as long as petals; inner stamens connate to V 3 of their length; 
anthers rounded; pistil with short style; hypogynous scales emarginate, 
subquadrate, ca. 0.5 mm long; follicles 7—10 mm long, dark red, the style 
filiform, reflexed, short, persistent; fruit with persistent perianth lobes; 
seeds brown, 2 mm long, lanceolate. June— July. 

Alpine zone, rocks, stony slopes, near glaciers, on ancient moraines, 
pebbly lichen tundras, near streams on moistened sites.— W.Siberia, 
Alt. Gen. distr. : Mong. (NW Mong.). Described from Altai. Type in 
Leningrad. 

14. R. komarovii A. Bor. sp. nova in Addenda, VIII, p. 362.— Sedum 
polytrichoides Kom., non Hemsl. in Kom. and Alis., Opr. rast. 
Dal'nevost.kraya I (1931) 601. 

Perennial; roots slender; caudex long, slender, 0.5— 0.7 cm broad, 
covered in its upper part with acute, scalelike leaves, 2—3 mm long, 2—3 mm 
broad at base; stems 3—4, glabrous, densely leafy, ca. 10 cm long, 1 mm 
thick; leaves alternate, approximate, linear, subterete, fleshy, broader in 
upper part, entire, sometimes with 1 or 2 teeth, subobtuse, 1—2 cm long, 
1—2 mm broad, green; inflorescence leafy, ca. 1 cm in diamter, 1—5 -flowered, 
corymbiform, terminal, with 2—4 branches; pedicels shorter than flowers; 
flowers bisexual, sometimes pistils obsolete, 4 (5)-merous, 4—5 mm long; 
sepals narrow, linear, acuminate, 1.5—2 mm long, green, Y 2 — 2 / 3 times as 
long as petals; petals lanceolate -elliptic to lanceolate, subobtuse, pale 
yellow, persistent in fruit; epipetalous stamens shorter than petals, the 
episepalous as long as petals; hypogynous scales broader than long; 
follicles 7 mm long, lanceolate, the 2 mm style reflexed; seeds dark 
brown, nearly black, oblong, ca. 2 mm long, acute. June — July. 

Rocky slopes, crevices of moist, shady rocks, bluffs, at more than 
1,000 m.— Far East: Uss. (Khualaza volcano, collected by T. Shishkin). 
Endemic. Probably occurs in the adjacent part of Manchuria. Described 
from the Far East (Khualaza in the Suchan district). Type in Leningrad. 

Note. Closely related to the Himalayan S. crassipes Wallich, from 
which it is distinguished by larger, more robust caudex and stem, entire or 
subentire leaves, few-flowered inflorescence, shape and size of leaves; it 
has been confused with Sedum polytrichoides Hemsl., section 
G e nu i na, which it resembles in habitus, though differing sharply by the 
presence of a caudex with scalelike leaves. 



31 



39 



Section 3. CHAMAE-RHODIOLA Schrenk in Enum. pi. nov. I (l84l) 69.- 
Caudex thick, surrounded by a mass of persistent stems forming a tuft; 
stems numerous, slender, 1—2 mm in diameter; follicles connate at base. 



40 



Series 1. Integerrimae A. Bor. — Leaves linear to oblong, entire. 

15. R. quadrifida (Pall.) Fisch. et Mey. in Schrenk, Enum. pi. nov. I (1841) 
69.-Sedum quadrifidum Pall., Reise III (l 776) Anh. 730; Ldb., Fl. 
Ross. II, 177; Fedtsch. O. et B., Consp. Fl. Turk. 3, 71; Berger in Engl. u. 
Pr., Nat. Pflzfm. 18a (1930) 442.- Chamaerhodiola quadrifida 
Nakai, Rep. of the first Scient. exped. to Manchoukuo sect. IV, I, (1934) 30.— 
Ic: Pallas, 1. c.,tab. P., f. 1A.; Frod. in Acta Horti Gothoburg. V (1930) 

pi. H, textfig. 43-47. 

Perennial; root long, reddish, thick; caudex 1—3 cm thick, multicipital, 
black-brown, covered with scalelike, triangular, brown leaves 4 mm broad, 
4 mm long and with persistent old stems; stems slender, 0.5—1 mm thick, 
in age often turning black or red, capilliform, numerous, 3—10 (l5)cm high, 
densely leafy, more or less erect, strict; leaves alternate, sessile, linear, 
subterete, fleshy, entire, acute, 5—8 (l2)mm long, 1 mm broad, usually erect; 
inflorescence few-flowered, corymbiform, ca. 1 cm in diameter; flowers 
dioecious, usually 4-merous, small; as long as or longer than pedicels; 
sepals 3 mm long, linear -lanceolate, obtuse, greenish; petals oblong -obovate, 
yellow, sometimes reddening at the apex, often drying green, obtuse, ca. 4 mm 
long; stamens 8, as long as or slightly longer than petals, with yellow 
filaments and anthers; hypogynous scales lanceolate -oblong, emarginate, 
red, nearly half the length of petals; follicles ca. 5 mm long, lanceolate, 
erect, the short beak slightly reflexed or erect, dark red at maturity; 
seeds 2 mm long, alate, brown, oblong. May— June. 

Alpine zone, less often in Polar-Arctic regions, pebbly or mossy lichen- 
tundras, rocks, stony slopes, taluses near glaciers.— Arctic: Arc. Sib., 
(Urals), Arc. Eur.; W.Siberia: Alt.; E. Sib. : Dau., Ang. -Say., Lena-Kol. 
Gen. distr. : Dzu.-Kash., Tannu-Ola, Mong. Described from the S. Urals 
and Mt. Sokhondo in Dauria. Cotype in Leningrad. 

Note. A form with reddish flowers occurs sometimes in the Altai. 
R. coccinea (Royle) A. Bor. was considered a synonym of R. quadrifida 
F. and M. The latter Central Asian species extends to the northeast 
as far as the Saur Range. 

16. R. kashgarica A. Bor. sp. nova in Addenda VIII, p. 362. 

Perennial; root slender, grayish, cordlike; caudex multicipital, branching, 
ca. 0.5—1 cm in diameter, with remnants of preceding year's stems and 
scalelike, triangular, acuminate leaves, 2—5 mm long, 3—5 mm broad; stems 
reflexed, divergent, numerous, 3—5 (l0)cm high, 0.5—1 mm in diameter, the 
old stems grayish; leaves alternate, subhorizontally spreading, oblong or 
linear -lanceolate, slightly narrowing toward base, sessile, fleshy, 3—10 mm 
long, 1—2 mm broad, entire, subobtuse; inflorescence few -flowered, compact 
or rather loose, corymbiform or multicipital, 0.5— 1 cm in diameter, 
0.4—0.6 cm high; flowers subsessile or pedicels shorter than flowers, 
elongating in fruit; flowers mostly 4-, rarely 5 -merous, dioecious, golden- 
yellow, ca. 3 mm long; sepals linear, acute, 2 / 3 as long as or subequal to 



32 



petals; petals 3—4 mm long, oblong-lanceolate, constricted at the apex, 
subobtuse; stamens slightly shorter than to slightly exceeding petals, 
with yellow filaments and rounded yellow anthers; pistils obsolete in 
staminate flowers; hypogynous scales subquadrate or more elongated; 
follicles 3—4 mm long, ovoid, with short, reflexed beak; seeds ca. 1.5 mm 
long, lanceolate, brown. June— July. (Plate III, Figure 5a). Rock streams, 
rocks, and old moraines in the high -mountain zone. — Centr. Asia: T. Sh. 
(the part of Lake Chatyr-Kul adjacent to Kashgaria, Aksai River). 
Gen. distr.: Dzu. -Kash. (Kashgar). Described from the Billuli River in 
Kashgaria. Type in Leningrad. 

17. R. pamiroalaica A. Bor. sp. nova in Addenda VIII, p. 363. 
Perennial; root robust, thick; caudex robust, woody, 1.5—3 cm broad, 

with remnants of preceding year's stems and scalelike, membranous, 
triangular-lanceolate, acuminate leaves (4)6— 8 mm long, (l . 5)3— 6 mm 
broad; stems curved, numerous, 10—20 (30) cm long, 2 mm in diameter, 
sulcate in lower part; leaves alternate, remote, linear, linear -lanceolate to 
lanceolate, broadening toward base, sessile, fleshy, 0.7—1.5 cm long, 
1.5—2 mm broad, entire, acuminate; inflorescence many -flowered, dense, 
less often loose or few-flowered, corymbiform -paniculate, 0.5— 1 cm 
long, 1 — 1.5 (2) cm in diameter, bracteate; flowers 5 -sometimes 6-merous, 
dioecious, pale yellow, ca. 4 mm long; as long as or longer than pedicels; 
sepals lanceolate or linear, subobtuse, 2 mm long, greenish yellow; petals 
subobtuse, lanceolate or linear, 4 mm long, pale yellow; stamens shorter 
than petals, filaments pale yellow; anthers rounded, yellow; hypogynous 
scales quadrate, entire at the tip or slightly emarginate; follicles 
ca. 4 (6) mm long, oblong, with straight filiform beak, ca. 1 mm long; in 
mature fruit beak sometimes reflexed; seeds 2 mm long, 0.5 mm broad, 
lanceolate, brown. Fl. June — July, fr. June — August. (Plate III, Figure 6a— b). 

Stony soils, pebbles in the alpine zone.— Centr. Asia: T. Sh. (district 
of Karakol), Pam. -Al. (Shugnan, Darvaz, Pamir, Alai Range). Gen. distr. : 
Dzu. -Kash. (Kuldja, Kashgaria), W. Mong. Described from the Vanch 
River in Darvaz. Type in Leningrad. 

18. R. coccinea (Royle) A. Bor. comb. nov. — Sedum coccineum Royle, 
Illustr. Bot. Himal. I (1839) 223.- S.quadr ifidum auct.-Ic: Royle, 
I.e., tab. 48, f. 3. 

Perennial; root long, thick; caudex thick, branching, mult icipital, gray - 
brown, covered with scalelike, rounded -triangular, brown leaves 6— 8 mm 
broad, ca. 5 mm long, and with remnants of old stems; stems slender, 
1.5—2 mm thick, not blackening in age, not capillaceous, 5—15 cm high, 
densely leafy, erect or curved; leaves alternate, sessile, lanceolate, 
fleshy, entire, subobtuse, 5—6 mm long, 1—1.5 mm broad; inflorescence 
corymbiform, compact, few-flowered, 0.8—1 cm in diameter; flowers 
dioecious, ca. 4 mm long, usually 5 (4)-merous, longer than pedicels; sepals 
2 / 3 as long as corolla, oblong, subobtuse, red; petals oblong-ovate red, 
obtuse, constricted at the apex, ca. 4 mm long; stamens 10 (8), shorter 
than petals, with rounded yellow anthers and red filaments; hypogynous 
scales oblong, a / 3 as long as petals; follicles ovoid or oblong-ovoid, red, 



33 



with thick, very short reflexed beak; seeds 1 — 1.5 mm long, brown, oblong. 
June — July. 

Alpine belt, stony soils, rocks, at up to 2,000 m. — Centr. Asia: T. Sh., 
Dzu. -Tarb. (Saur Range, Muztau Range), Pam. -Al. Gen. distr. : Dzu.-Kash. 
(Kuldja, Kashmir). Described from Kashmir. Type in London. 

Note. See note referring to R. quad r if id a (Pall.) F. et M. 



Series 2. Oblongae A. Bor. Leaves oval or oblong-elliptic, denticulate 
or dentate or subentire. 

19. R. gelida Schrenk, Enum. pi. nov. I (1841 ) 67. — S e d um gelid um 
Ldb., Fl. Ross. II (1844-1846) 177; Fedtsch. O. et B., Consp. Fl. turk. 3 (1909) 
71; Kar. et Kir., Enum. pi. Song, (l 842), No. 346. - C h a m a e r ho d i o 1 a 
gelida Nakai in Rep. first Scient. exped. Manchoukuo sect. IV, I (1934) 30.— 
Ic: Frod. in Acta Horti Gothoburg. V (1930) textfig. 76-86. 

Perennial; root robust, vertical; caudex thick, multicipital, 2—7 cm long, 
0.5—0.8 cm broad, covered with scalelike, acuminate, triangular -ovate leaves 
in upper part; leaves 0.5—0.7 cm long, 0.4 cm broad at base; stems 
numerous, those from the preceding year persisting on the caudex, 
3—5 (l0)cm high, slender, 1 mm in diameter, curved; leaves alternate, flat, 
.~ ovate -oblong, denticulate or subentire, 0.8—1 cm long, 2—5 mm broad near 
middle of leaf; inflorescence 1—2 cm in diameter, 1 — 1.5 cm long, usually 
many -flowered, dense; flowers dioecious, usually 4-, less often 5-merous, 
4—5 mm long, longer than the short, thick pedicels; sepals lanceolate or 
oblong-elliptic, subobtuse, 3 mm long, yellow; petals obtuse, lanceolate or 
oblong-elliptic, 4 mm long, yellow; stamens 8, less often 10, slightly longer 
than petals, their yellow filaments adnate to petals through much of their 
length; ovary with a recurved style at the tip; stigma disk-shaped; 
hypogynous scales elongated, 3 times as long as broad, yellow, mostly 
emarginate; follicles 4—5 mm long, reddish, with divergent short beak; 
seeds ca. 1.5 mm long, lanceolate, brown. Fl. June— July, fr. July— September. 

Alpine zone, stony -pebbly rock streams, rock crevices, alpine tundra. — 
Centr. Asia: T. Sh. (often), Dzu. -Tarb. (more rarely), Pam. -Al. (Alai and 
Trans -Alai ranges). Gen. distr.: Mong. (W.). Described fromDzhil- 
Karagai, Dzungaria. Type in Leningrad. 

20. R. recticaulis A. Bor. sp. nova in Addenda VIII, p. 364. 
Perennial; root thick, woody; caudex robust, multicipital, 3—4 (6) cm 

broad, with separate branches, ca. 1.5 cm broad, covered in its upper part 
with brownish, membranous, scalelike, triangular, obtuse leaves ca. 1 cm 
long and 1 cm broad, the upper scalelike leaves longer and the lower 
broader; stems numerous, those from the preceding year persisting, 
(8) 1 2—15 cm high, 1.5—2 mm in diameter, mostly erect, slightly sulcate, 
leafy; leaves alternate, oval to oval-oblong, coarsely dentate, acuminate, 
0.8—1 cm long, 0.2—0.3 cm broad, erect, dark green; inflorescence dense, 
compact, many -flowered, more rarely few-flowered, corymbiform, capitate, 
ca. 1.5—2 cm in diameter, leafy; flowers dioecious, 4 -merous, small, 
longer than pedicels; sepals half as long as petals, oval, obtuse, reddish or 
red; petals oblong-elliptic, obtuse, 4 mm long, yellow; stamens longer than 



34 




PLATE III. 1-Rhodiola arctica A.Bor.: a) fruit, b) flower; 2 - R. sa chal i nensis A. Bor.: fruit, 

b) flower; 3 - R.he t er od on ta (Hook, et Thorns.) A. Bor.: a) flower; 4 - R. 1 i tv i no v i i A. Bor.: 

a) fruit; 5- R.kaschgarica A.Bor.: a) flower; 6 - R. pa mi ro a 1 a i c a A. Bor.: a) flower, b) fruit. 



35 



45 



46 



petals, with yellow filaments and rounded anthers; hypogynous scales 
subquadrate, nonemarginate; pistils with disk-shaped stigma; follicles with 
a short beak; seeds 2 mm long, 0.5 mm broad, oblong, brown. Fl. June — 
August, fr. July — September. 

Alpine meadows in the high -mountain alpine zone.— Centr. Asia: 
Pam.-Al. (often in Darvaz, Shugnan, Vakhan, Pamir, less often in Trans - 
Alai and Alai ranges), T. Sh. (in the part adjacent to Kashgaria, district 
of Karakol). Gen. distr. : Iran. (Afghanistan), Dzu.-Kash. Described from 
the Koitezek Pass in the Pamir. Type in Leningrad. 

Note. The Pamir -Alai species is closely related to R. gelida 
Schrenk, from which it is distinguished by its much larger scalelike 
leaves, red sepals, robust, woody caudex, greater height, erect, more robust 
stems, and shape of hypogynous scales. The Tien Shan species R. gelida 
Schrenk penetrates into the Pamir -Alai only from the north; R. recticaulis 
A. Bor. penetrates into Tien Shan only in its eastern part (district of 
Karakol). 

21. R. litvinovii A. Bor. sp. nova in Addenda VIII, p. 3 64. 

Perennial; root thick, ca. 30 cm long and ca. 2 cm broad in its upper part; 
caudex robust, multicipital, 3—4 cm long, ca. 1—2 cm broad, with few old 
stems, covered in its upper part with triangular, ovate, light yellow, 
membranous, large, acuminate leaves 1—1.5 cm long, 1 — 1.5 cm broad at 
base, the upper, scalelike leaves longer, the lower short, broad, 
semiorbicular; stems numerous, 10 — 17 cm long, slightly sulcate, erect, 
rather densely leafy, 2—4 mm in diameter; leaves alternate, flat, elliptic, 
with obtuse, unequal, deep teeth, entire only toward base, with cuneate, 
petiolate base, subobtuse, 1 — 1.5 cm long, 0.3—0.5 cm broad, all leaves erect, 
pale green, always dryingyellow-green; inflorescence dense, many -flowered, 
compact, 2—2.5 cm in diameter, 1 — 1.5 cm long, leafy; flowers yellow, 
4 - or 5 -merous, dioecious, small, ca. 4 mm long, as long as or longer than 
pedicels; stamens longer than petals, with yellow filaments and rounded 
yellow anthers; hypogynous scales quadrate, entire at the apex; follicles 
ca. 8 mm long, gradually passing into a long filiform beak 1—1.5 mm long; 
seeds 1.5—2 mm long, 0.5 mm broad, oblong-lanceolate, brown. Fl. June- 
July, fr. June— August. (Plate III, Figure 4a). 

Stony and pebbly soils, mountain passes, near glaciers in the alpine 
zone.- Centr. Asia: T. Sh., Pam. -Al. Gen. distr. : Mong. (N. and W.)., 
Dzu.-Kash. Described from the Kenkol Pass in the Andizhan District. 
Type in Leningrad. 



Genus 6 99. SEDUM * L. 
L. Gen., pi. ed. 5 (1754) 197. 

Flowers yellow, white, pink, red, rarely sky-blue, mostly 5 -merous, more 
rarely (4) 6— 9 -merous; petals connate only at base or free; stamens twice 
as many or of same number as petals; stamens opposite to petals usually adnate 
to petals at base; fruit an aggregate follicle; follicles as many as petals, 

From the Latin sedare.to pacify - the succulent leaves are used as an analgesic for wounds - or from 
sedere.to sit - many species are prostrate. 



36 



many-seeded, with short beak. Annuals and perennials of various aspect. 
Most species of the Sedum genus are distributed throughout the Temperate 
Zone of the northern hemisphere, mainly in Eurasia; Sedum is most 
widely distributed in the Himalayas, China, and Japan. The eastern and 
southeastern parts of N. Am. contain more species than the western part; 
only single species occur in the southern hemisphere (Centr. Afr., 
Madagascar, S. Am.). 

1 . Perennials with thick, short caudex or with tuberous, fusiform and 
woody roots or with creeping rhizome 2 - 

+ Annuals, less often biennials, with slender roots; leaves alternate, 

mostly semiterete or terete, less often rather flat, linear or 
lanceolate, rarely to oblong -ovate. (Section 5. Epeteium 

Boiss.) 5 - 

2. Under shrubs with woody roots and stems lignifying from base; 
leaves flat, ovate, petiolate, coarsely and unequally dentate. 

(Section 2. Populisedum Berger) 16. S. populifolium Pall. 

+ Herbs, not usually with woody roots, their above-ground parts 

decaying annually, or else evergreen 3 - 

3. Plants with creeping rhizome, usually evergeen, cespitose at base, 
sometimes with white globose underground buds; stem creeping, 
ascending or erect, branching from base; leaves semiterete or 
scalelike, thickened or flat, spatulate, but then with white, pinkish, 

or reddish flowers. (Section 4. Eu sedum Boiss.) 17. 

+ Plants with thickened roots or with short rhizome and tuft of 
slender roots or with long creeping rhizome, but then flowers 
yellow; stems erect or ascending, not branching, less often 
subprostrate; leaves flat, ovate to lance-linear 4. 

4. Flowers yellow; roots not tuberous; rhizome short or else long, 
creeping, rooting, but then plants evergreen; leaves oblong-oval to 
linear or lanceolate. (Section 3. Aizoon Koch) 34. 

+ Flowers greenish, white, pink, or red; roots thick, tuberous, fusiform, 
or else rhizome short, inconspicuous, with tuft of slender roots; 
leaves rounded to oblong -elongate. (Section 1. Telephium 
S.F.Gray) - • ■ •• 40 ' 

5 (l). Flowers 4- or 5-merous; stamen 4-5, sometimes also 4 (5) 

R 

obsolete 

+ Flowers 5 - or 6 -merous, less often 7-9-merous; stamens 10-12 

(14-18) 9 - 

f 6. Calyx exceeding corolla '• 

+ Calyx shorter than corolla 8 - 

7. Sepals glabrous, linear -terete or oblong-ovate; leaves glabrous 

or sparsely ciliate -margined. (Caucasus, Central Asia) 

S. tetramerum Trautv. 

+ Sepals ciliate, ovate; leaves ciliate. (Eur., Crim.) . 

54. S. aetnense Tin. 

8. Subglabrous plants 2—6 cm high; leaves broadly elliptic to ovate, 
imbricate; calyx % — 1 / 2 as long as petals, glandular -pubescent; 
flowers 4, rarely 5-merous 52. S. rubrum (L.) Thell. 

+ Glandular -pubescent, succulent, glabrous reddish plants 5— 15 cm 
high; leaves oblong-linear, semiterete; calyx V 4 — V 3 as long as 
petals; flowers 5-merous 51. S. rubens L. 



37 



48 



9. Pedicels longer than flowers 10. 

+ Pedicels shorter than or as long as flowers or else flowers 

sessile 11. 

10. Glandular -pubescent plants; petals pink 48. S. villosum L. 

+ Glabrous plants; petals yellow 50. S. nanum Boiss. 

11. Flowers yellow. Arcto-alpine species 49. S. annuum L. 

+ Flowers white, pinkish, reddish, or greenish 12. 

12. Glabrous, reddish-brown plants; corolla greenish white or greenish 
pink. (Europe) . . . 47. S. atratum L. 

+ Glandular-pubescent, green plants, sometimes reddening; corolla 

white, pink, or greenish 13. 

13. Flowers on pedicels 3— 4 mm long; follicles connate nearly to the 

middle; inflorescence a regular corymb 

46. S. corymbosum Grossh. 

+ Flowers subsessile; follicles connate at base; inflorescence 

corymbiform 14. 

14. Follicles stellately spreading; flowers white or pinkish; petals 
lanceolate, stellately -divaricate 15. 

+ Follicles directed upward, the long beak 1.5 times as long as fruit; 
flowers pink, 5-merous; petals narrowly lanceolate, usually 
erect 45. S. pallidum M. B. 

15. Flowers 5-merous; plants not branching from base, stem erect; 
petals 5—6 (7)mm long; follicles 5— 6 mm long, lanceolate, with a 
protuberance at base ventrally. (Caucasus, Mtn. Turkm., Iran) .... 
43. S. pentapetalum A. Bor. 

+ Flowers 6 (7— 9)-merous; petals 4— 5 mm long; follicles 2— 4 mm 
long, gradually tapering toward base, without conspicuous 
protuberance at base 16. 

16. Plants branching from base, with numerous ascending stems; 
follicles 2— 3 mm long, rounded -ovoid or ovoid; flowers 6 (7—9)- 
merous. (Eur. Caucasus) 42. L. hispanicum L. 

+ Plants with ascending solitary stem not branching from base; 

follicles 4 mm long, oblong -triangular, gradually broadening toward 
base; flowers 6-merous. (Pam.-Al.). . . . 44. S. bucharicum A. Bor. 

17. (3). Flowers yellow 18. 

+ Flowers white, pink, or red 21. 

18. Leaves ovate, convex on the back, obtuse, without basal appendage; 
flowering shoots 5— 15cm high 40. Caere L. 

+ Leaves linear or lance-linear, terete or semiterete, with or without 

basal appendage; flowering shoots 8—2 5 (40)cm high 19. 

19. Leaves with basal appendage; follicles erect; fruiting shoots erect, 
much longer than the sterile, scarcely branching. Europe 2 0. 

+ Leaves without basal appendage; follicles substellately spreading; 

fruiting shoots scarcely exceeding or about as long as the sterile, 

branching, curved, creeping. (E. Asia) 

41. S. polytrichoides Hemsl. 

20. Leaves linear, terete, ca. 0.5 cm long, 1 cm broad, subobtuse. Plants 
8— 18 cm high 39. S. sexangulare L. 

+ Leaves linear -subulate, semiterete, ca. 1.5—2 cm long, 2— 3 mm broad, 
acute. Plants 15-25 (40)cm high 38. S. reflexum. 



38 



21. Leaves opposite, flat, broad, oval or obovate, spatulate. (Subsection 

1. Spathulata A. Bor.) 22. 

+ Leaves alternate, terete or thickened, linear- subulate to oblong. 

(Subsection 2. Crassi folia A. Bor.) 28. 

22. Plants with globose white underground buds. (Series P r op o nt i c a e 
Berger) 23. 

+ Plants without white underground buds. (Series Involucratae 

Maxim.) 24. 

Leaves 6— 30 mm long, 5 — 18 mm broad; inflorescence branches 
villous -scabrous 29. S. obtusifolium C.A.M. 

+ Leaves 15— 40 mm long, 10 — 30 mm broad; inflorescence branches 

glandular -pubescent ...30. S. listoniae Vis. 

24. Small plants 3—10 cm high; leaves 2—6 mm long, 1—4 mm broad, 

entire, glabrous, spatulate -ovate; flowers 5— 7 mm long 

24. S. stevenianum Rouy et Camus. 

+ Larger plants 5— 20 (25) cm high; leaves 8— 2 5 mm long, 5— 16 mm 
broad, broadly dentate or crenate or obscurely emarginate, 
petiolate 25. 

25. Follicles stellately spreading, with two ventral protuber- 
ances; flowers 5—8 mm long 25. S. stoloniferum Gmel. 

+ Follicles not stellately spreading, without umbo ventral protuber- 
ance; flowers (7)10— 15 mm long 26. 

26. Inflorescence dense, compact, surrounded by large terminal 
leaves surpassing the inflorescence; corolla 1.5 times as long 

as calyx 28. S. involucratum M. B. 

+ Inflorescence not surrounded by terminal leaves; corolla twice as 

long as calyx 27. 

27. Corolla pink or purple; filaments red, anthers orange -red; petals 
lanceolate, acute; fruit reddish 26. S. spurium M. B. 

+ Corolla white or pale yellow, cream-colored; filaments white or 
yellow, later darkening; petals linear -lanceolate, acuminate; fruit 
green 27. S. oppositifolium Sims. 

28. Corolla red or pink, sometimes whitish; flowering shoots 4—10 cm 
high; corolla 1.5—2 times as long as calyx 29. 

+ Corolla white, rarely pinkish; flowering shoots (5) 10— 20 (30) cm 

high; corolla 2—4 times as long as calyx 30. 

29. Corolla red, sometimes whitish; inflorescence corymbiform or 
paniculate, rather loose, few -flowered; leaves oblong, 3—5 mm long, 
1-1.5 mm broad 31. S. tenellum M. B. 

+ Corolla pink, sometimes whitish; inflorescence capitate, dense, 

many -flowered; leaves narrowly linear or lanceolate, 4— 6 (8) mm 
long, ca. 1 mm broad *32. S. lydium Boiss. 

30. Leaves oblong or oblong -ovate, 7—10 mm long, 1-^-2 mm broad; 
sepals obtuse, V 4 — V 3 as long as corolla , . 33.. S. album L. 

+ Leaves linear-subulate to oblong -lanceolate; corolla 2—3 times as 

long as sepals 31. 

31. Leaves obtuse, glabrous 32. 

+ Leaves acuminate or if obtuse then minutely tuberculate on the 

surface 33. 



39 



51 



32. Flowering shoots 4— 6 cm long; inflorescence many -flowered; 
corolla 3— 4 mm long 34. S. gracile C. A. M. 

+ Flowering shoots (5)10—15 (20) cm; inflorescence few-flowered; 

corolla 6 — 7 mm long 35. S. lenkorianicum Grossh 

33. Leaves acuminate, glabrous, linear -subulate; sepals subacute. 
(Caucasus, Asia Minor) 36. S. subulatum (C.A. M.) Boiss. 

+ Leaves obtuse, minutely tuberculate on the surface, linear or 

linear -oblong; sepals subobtuse. (Central Asia) . . 37. S.Alberti Rgl, 

34. (4). Entire plant densely pubescent .... 23. S. selskianum Rgl. et Maak. 
+ Glabrous plants. (Series Gl a b r a e A. Bor.) 35. 

35. Plants with short thick rhizome; stems 25—80 cm long, strong, 
erect; inflorescence surrounded by terminal leaves 36. 

+ Plants with elongated creeping branching rhizome, mostly weak, 

ascending or erect 37. 

36. Stems 25—45 cm high, few, leaves subacute, elongate, lanceolate to 
linear, 5— 8 cm long, 1—2 cm broad 17. S. aizoon L. 

+ Stems to 85 cm long, solitary, with obtuse, ovate -lanceolate leaves 

8— 10 cm long, 3— 4 cm broad 18. S. hyperaizoon Kom. 

37. Leaves in whorls of 3, ovate or obovate -lanceolate, subobtuse, 

broadly serrate -dentate, with adjacent uncinate teeth 

22. S. litorale Kom. 

+ Leaves alternate, linear to elongate -lanceolate and spatulate- 

elliptic, crenate or coarsely and obtusely dentate 38. 

38. Leaves linear, coarsely and sparsely dentate at the apex; petals 
about twice as long as stamens. Shrubby, many- stemmed plants 
20. S. middendorfianum Maxim. 

+ Leaves oblong-elliptic or spatulate -elliptic, sometimes elongate, 
obtusely serrate; stamens as long as or slightly shorter than 
petals 39. 

39. Stamens slightly shorter than petals. Ascending plants, rooting 
and prostrate, many-stemmed, cespitose; stems evergreen, 

persistent stem; flowers yellow; sepals lanceolate, equal 

21. S.hybridum L 

+ Stamens as long as petals. Erect plants with few stems; flowers 

orange -yellow with broadened ovate base and elongated apex 

19. S. kamtczaticum Fisch. 

40.(4). Tall plants, (15) 30-70 (100) cm high with strong stem; leaves 
3—10 cm long; roots tuberous or rhizome very short, scarcely 
developed, with tuft of slender roots. (Subsection 1 . Erecticaulia 

Praeger) 43. 

+ Low plants, 5—15 (25) cm high, the weak stem creeping or ascending; 
leaves 0.5—2.5 cm long; rhizome branching, cespitose, with 
numerous fruiting and sterile shoots. (Subsection 2. Humili - 
caulia Praeger) 41.. 

41. Plants 5— 7 cm high; flowers blue or purple; stamens shorter than 

or as long as petals; leaves ca. 1 cm long, from 2—3 to 5—7 mm 

broad, alternate or opposite, connate at base 42. 

+ Plants 10— 20 (25) cm high; flowers pink, light purple -lilac; 

stamens slightly longer than petals; leaves 1.5—2 cm long and 

about as broad, opposite, with cordate base 13. S. ewersii Ldb 



40 



42. Leaves opposite, connate at base, broadly ovate or rounded, 
0.7—1 cm long, 0.5—0.7 cm broad, obtuse, without red dots; sepals 

1.5— 2 mm long, linear-subulate, acute. (Sakhalin) 

14. S. pluricaule Kudo. 

+ Leaves alternate, oblong-lanceolate to sublinear, ca. 1 cm long, 

0.2—0.3 cm broad, with red dots; sepals 3 mm long, oblong, acute. 

(E. Siberia) 15. C. cyaneum Rud. 

43. Caudex very short or obsolete, with a tuft of slender roots 44. 
+ Roots tuberous, rounded or fusiform. (Series E u - T e 1 e ph i a e 

A. Bor.) 49. 

44. Leaves in whorls of 4 or 5, the lower sometimes in pairs; fleshy 
white buds in leaf axils and on rhizome. (Series Viviparae 

A. Bor.) 45. 

+ . Leaves opposite, sometimes in threes, orbicular to broadly oval, 

cordate and ovate-cuneate or alternate, sometimes in pairs, 
approximate, elliptic, lanceolate or oblong -ovate with cuneate 
base; axillary buds absent. (Series Fa s c i c u 1 at a e A. Bor.) . . 46. 

45. Leaves longer than internodes, 5—8 cm long, 1.5—3 cm broad, often 
with black dots; flowers ca. 4— 5 mm long .... 7. S. verticillatum L. 

+ Leaves usually shorter than internodes, 3—4 (5) cm long, 1.2 — 1.5 cm 

broad, with black dots; flowers ca. 3 mm long 

8. S.viviparum Maxim. 

46. Leaves opposite or so'metimes approximate, alternate, orbicular or 
ovate 47. 

+ Leaves alternate, oblong -ovate or elliptic -lanceolate 48. 

47. Leaves orbicular or broadly oval, amplexicaul, sinuate-dentate, 

subobtuse; flowers purple, mostly 4 -merous 

9. S. ussuriense Kom. 

+ Leaves ovate with broadly cuneate base, broadly on the margin, 

subacute; flowers whitish or pinkish, 5 -merous 

12. S. alboroseum Baker. 

48. Stems 30— 6 0cm high; leaves 3— 7 cm long, 0.7— 2.5 cm broad; 
inflorescence usually 3—5 (9) cm in diameter, 2—5 (7) cm long; 

petals lanceolate 10. S. pallescens Freyn. 

+ Stems 60— 100 cm; robust; leaves 7— 10 cm long, 1 .5— 4 cm broad; 
inflorescence usually 7— 10 cm in diameter; 10— 30 cm long; petals 
broadly lanceolate 11. S. eupatorioides Kom. 

49. Leaves alternate 52. 

+ Leaves opposite 50. 

50. Leaves large, fleshy, cochlear -concave, obovate, deeply cordate at 
base, amplexicaul, broader than long, 5—7 cm broad, 3—5 cm long 
6. S. caucasicum (Grossh.) A. Bor. 

+ Leaves oval, orbicular, or oblong, amplexicaul or nonamplexicaul, 

2 — 13 cm long, 1—5 cm broad, usually longer than broad 51. 

51. Roots rounded -elongated or napiform; leaves orbicular or oval, 
broadening toward base, usually amplexicaul, 4—5 cm long, 2—3 cm 
broad; flowers 5— 6 mm long 5. S. telephium L. s. s. 

+ Roots fusiformly thickened; leaves oblong -elliptic, nonamplexicaul, 

5 — 13 cm long, 2— 5 cm broad; flowers 3— 4 mm long 

4. S. maximum Suter. 



41 



52. Leaves with cordate -amplexicaul base, entire or sometimes with 
1 or 2 teeth at base; inflorescence composite, racemose- 
paniculate 2. S. mugodsharicum A. Bor. 

+ Leaves with cuneate base, dentate; inflorescence corymbiform or 

corymbiform -paniculate 5 3. 

53. Flowers 4 -merous, bisexual, sometimes unisexual; stamens sessile; 

leaves dark green, unequally sinuate -dentate 

3. S. parvistaminex m V. Petrov. 

+ Flowers 5 -merous, always bisexual; stamens as long as or slightly 

longer than petals; leaves usually — mainly at the summit — 
_„ unequally dentate, or regularly acutely dentate along almost the 

whole margin 1. S. purpureum Schult. 



Section 1. TELEPHIUM* S. F. Gray, Nat. Arrang. Brit. PI. II (1821) 539. 
Flowers 5 -merous, white, pink, red, blue, or greenish-yellowish. Rhizome 
thick, reduced, without scalelike leaves, or else rhizome long, slender, 
rooting, with slender or robust fusiform or tuberous roots; stems erect 
or ascending, less often subprostrate, mostly annual; leaves flat, broad, 
fleshy. 

European and Asiatic species, occurring mainly in China, Japan, and 
N. America. 

Economic importance. Some species of the section Telephium are 
cultivated as ornamental plants (S. spectabile from Japan, S. a n a - 
c amp s e r o s), and several others may be recommended as ornamentals 
(S.ewersii, S. cyaneum). 



Subsection 1 . ERECTICAULIA Praeger in Journ. Hort. Soc. XLVI (.1 921 ) 
79, pro serie. — Stems strong, erect, simple, tall; leaves 5— 10 cm long. 



Series 1. Eu-Telephiae (Praeger) A. Bor.; Praeger, 1. c, p. p., pro 
group. — Roots tuberiform, tuberculate; leaves flat, large, broad, alternate 
or opposite; stems robust, (15) 30—80 cm high. 

1. S. purpureum (L.) Schult., Oestr. Fl. ed.2, I (1814) 686; Link, Enum. 
berol. (1821) I, 437; Ldb., Fl. Ross. II, 181; Shmal'g., Fl. I, 360; S.fabaria 
auct. — S. purpurascens Koch, Syn. ed.2 (1846) 284.— S. telephium 
/3 et 7 purpureum L., Sp. pi. (1753) 430; Berger in Engl. u. Pr. Nat. 
Pflzf. 18a (1930) 444.- S. telephium Kryl., Fl. Zap. Sib. VI (1931) 1409; 
FrOd. in Acta Horti Gothoburg. V (1930) 61 (subsp. pu r pur eum). — 
S. vulgare Link, 1. c. (1821)437 p. p.— S. maritimum Bohusl. in Rupr., 
Fl. Samoj.II (1845) 10, 34; Erman, Archiv IV (1848) 66.- S. bohuslavii 
Rupr., Beitr. Pfl. Russ. Reich. VII (1850) 1 8. - S. a r c ha ng e 1 i c u m Rupr. 
nomen in sched. - Ic: Rchb., Ic. Fl. Germ. XXIII (1898-1899) tab. 44; 
Praeger in Journ. Hort. Soc. (l92l) f. 36, 38; Borisova in Sorn. rast. 
SSSR, III (1934) f. 238, 3 (seed).— Vernacular name: zayachya kapusta; 
Kirgiz — Koko-maral, Turkic — kantibaer, Kalmyk — uden'edszuun, 
tolstolistnik, skripun, kotovik, tselistnik, saigach'ya kapusta, etc. 

* From the Greek telephion or telephonion, the name of a thick-leaved, white -flowering plant, 
according to Dioscorides and Pliny. Later this name was given to a genus of the family Caryophyllaceae. 

42 



Perennial; roots tuberiform, tuberculate; stems erect, solitary or few, 
(15) 30—60 cm high; leaves alternate, unequally toothed mainly at the apex, 
ovate-oblong or oblong, the upper sessile, the lower cuneate -tapering, 
acuminate or subobtuse, 2—7 cm long, 1 — 3 cm broad, dark green; 
inflorescence dense, corymbiform; flowers 5-merous; calyx ca.2.5mm 
long, with lanceolate, acute lobes, green, connate at base; petals 2—3 times 
as long as calyx, elliptic -lanceolate, acute, ca. 5—6 mm long, reflexed from 
the middle, purple; stamens opposite to petals, united l L— 1 / 2 above base, 
about as long as petals; hypogynous scales yellow, 0.5 mm long, linear- 
oblong, emarginate; follicles erect, red or pink, ca. 6 mm long, with short, 
slightly reflexed beak, shorter than stamens; seeds small, numerous, to 
1 mm long, brownish, tapering to one end. Fl. July and beginning of August, 
two weeks earlier than S. maximum s. lat. 

Fields, plowland, sometimes cultivated fields, roadsides, as a weed, in 
water meadows, clearings, among shrubs, on rocky and stony argillaceous 
slopes.— European part : Kar. -Lap., Dv. -Pech., U. Dnp., U. V., V. -Kama, 
M. Dnp., V.-Don, BL, L. Don, L. V., Crim.; W.Siberia: Ob, U. Tob., Alt., 
Irt.; E.Siberia: Yenis., Ang. -Say., Dau., Lena -Kol.; Far East: Ze.-Bu., 
Uss., Uda, Okh., Kamch., Sakh. Gen. distr.: almost all of Europe and 
Mong. (Tuva ASSR), Jap.-Ch. (Manchuria, N. Korea, Japan, China), N. Am. 
(introduced). Described from Europe. Type in London. 

Note. Within its wide distribution area, S. pu rp u r eu m varies greatly 
in aspect; it is a highly polymorphic species. Var. p lu r ic au 1 e Maxim., 
distributed in E. Siberia, is a lower plant with many stems, leaves subentire, 
petals purple, 4— 5 mm long (Kamch., Kirensk, Sakh.). S.maritimum 
Bohusl. (S. bohuslavii Rupr.), described from Dv. -Pech. and V.-Kama 
(Perm) is distinguished from the type by its bright pink flowers and ovate - 
rhomboid, coarsely dentate, light-colored leaves. 

S.purpureum (L.) Schult., S. mug o d s h a r i c um A. Bor., S. p a r v i - 
stamineum V. Petrov, and a number of other species which do not occur 
in the Soviet Union, as for instance S. f ab a r ia Koch, can be united in a 
group of species characterized by alternate leaves and red or pink flowers. 

2. S. mugodsharicum A. Bor. sp. nova in Addenda VIII, p. 3 65. 
S.purpureum auct. 

Perennial; roots tuberous, fusiform, fascicular; stems erect, 35—60 cm 
high, smooth, pinkish, densely leafy; leaves alternate, acute, oblong, 2—4 cm 
long, 0.8—1.5 cm broad, entire, sometimes with 1 or 2 long narrow teeth 
in lower part, acute, broadening toward base, cordate -amplexicaul; 
inflorescence compound, racemose -paniculate, with short lateral branches 
arising in axils of upper leaves, many -flowered, 4—8 cm in diameter, 
7—10 cm long; flowers on pedicels shorter or as long as flowers; corolla 
twice as long as calyx; sepals linear-filiform, acuminate, green; petals 5, 
acute, elliptic or ovate, 5 mm long; stamens 10, those opposite to petals united 
with corolla to 1 / 3 , shorter than petals, those opposite to sepals as long as or 
slightly shorter than petals; pistils with long filiform style, convex dorsally, 
ovate; ovary stalked; follicles with reflexed beak; mature fruit unknown. 
July. 

Moist sites, mountain stream banks.— Centr. Asia: Ar.-Casp. 
(Mugodzhar Hills). Endemic. Described from the Mugodzhar Hills. 
Type in Leningrad. 



43 



56 



3. S. parvistamineum V. Petrov. in Bull. Jard. Bot. Princ. XXVI (1927) 
183. 

Perennial; stems 40—60 cm high; leaves alternate, dark green, unequally 
sinuate-dentate, oblong to ovate-oblong; inflorescence loose, branching- 
paniculate, with lateral branches arising from axils of upper leaves; 
flowers usually 4-merous, pale pink (?), sometimes unisexual; sepals 
twice as long as petals; stamens subsessile. Fl. from end of July to 
first frosts. 

Note. The author of this species presumes it to be native to the 
Far East, although there are no herbarium specimens from there. It 
flowers well and bears fruit in the Leningrad Botanical Institute of the 
Academy of Sciences. Unlikely to be a Far Eastern species, it may be 
of cultivated origin. 

4. S. maximum (L.) Suter, Fl. Helv. I (1802) 270; Ldb., Fl. Ross. II, 180, 
p.p.; Shmal'g., Fl. I, 360 p. p.; Kryl., Fl. Zap. Sib. VI, 1406 p. p.; Berger 

in Engl. u. Pr., Nat. Pflzfm. 18a (1930) 444 p. p.- S.telephium e 
maximum L., Sp. pi. ed. I (1753) 430 ; Frod. in Acta Horti Gothoburg. 
V (1930) 6 8.- S. lat ifolium Bertol., Amoenit. (1819) 366. - A na c a mp - 
seros maxima Haw., Syn. PL Succul. (l 819) 121 . — Ic. : Rchb., Ic. Fl. 
Germ. XXIII, tab. 45; Praeger in Journ. Hort. Soc. (1921 ) f. 36, 37.- 
Vernacular names: zayatch'ya kapusta, skripun, tolstolistnik. 

Perennial; roots thickened, fusiform, becoming gradually thinner; 
stems vigorous, erect, 40—80 cm high; leaves opposite, nonamplexicaul, 
oblong-elliptic, obtuse, mostly obscurely emarginate, dark green, 5 — 13 cm 
long, 2—5 cm broad; inflorescence broad, 6—10 cm in diameter, dense, 
corymbiform-paniculate, lower branches long, arising from axils of upper 
leaves; pedicels shorter than flowers; calyx with triangular, acute lobe, 
1 mm long, green, with short tube, rounded at base, 1 / 3 as long as corolla; 
petals ovate, acute, 3—4 mm long, ca. 1 mm broad, punctate, whitish pink; 
stamens 10, scarcely exserted from corolla, inserted at very base of 
petals, with rounded anthers; hypogynous scales linear, twice as long as 
broad; follicles erect, greenish, as long as petals, with linear beak; seeds 
oblong-ovoid, ca. 0.5 mm long, brown. July — October. 

Deciduous (broadleaf) forests.— European part : U. Dnp. (Mogilev), 
M. Dnp. (Kiev, Uman), BL, V. -Don (Kharkov); E. Siberia: Dau. (as a weed). 
Gen. distr.: Med., Bal. -As. Min., Centr. Eur., Scand., Atl. Eur. Described 
from Europe. Type in London. 

Note. S. maximum has been taken in a broad sense. In fact, 
there is a series of geographic races with varying ecology. The true 
S. maximum is typically west European, connected with broadleaf forests. 
S. maximum (L.) Suter and the following S. t e 1 ep hiu m (L.)s.s.and 
S. caucasisum A. Bor. can be united in a group of species characterized 
by opposite leaves, rarely by lower leaves alternate or approximate in 
threes, and by whitish, greenish, or yellowish, sometimes pinkish flowers. 

5. S. telephium L., Sp.pl. (1753) 430 s. s. - S. t elephium a album L., 
1. c. - S. vulgar e Link, Enum. h. berol. I (1821) 437, p. p.; Ldb., Fl.Ross. H, 
180, p. p.; Frod. in Acta Horti Gothoburg. V (1930) 67, p. p. - S. maximum 
auct. — S.polonicum Blocki in Deutsche Bot. Monatsschr. V (1887) 66. — 



5773 44 



Ic: Hegi, Fl. Mittel. Eur. IV, 2 (1925) 522, 523. - Exs. : PI. Finl. exs., No. 700; 
Fl. polon. exs., No. 633 (sub S. maximo Suter.). — Vernacular names: 
zayach'ya kapusta, skripun, tolstolistnik. 

Perennial; roots thickened, rounded -elongated, napiform; stems (10) 
20—40 cm high, usually arched -upcurved at base, mostly with shortened or 
scattered internodes; leaves opposite, orbicular or oval, broadening toward 
base, cordate, amplexicaul, strongly glaucous, drying dark green, densely 
disposed, obscurely emarginate, rarely dentate, (2)4—5 cm long, (l)2— 3 cm 
broad; inflorescence (3)5—10 cm in diameter, dense, corymbiform or loose, 
with elongated, scattered peduncles forming a corymbiform -paniculate 
inflorescence; lateral branches not always attaining top of inflorescence- 
pedicels shorter or longer than flowers; flowers ca. 5—6 mm or 2—3 mm 
long, broad; calyx rounded at base or subacute with triangular, lanceolate, 
acute sepals, V 4 — 1 / 3 as long as corolla; petals ovate, 1—2 mm broad, 5—6 or 
2—3 mm long, greenish, pale yellow, whitish-pinkish; stamens 10, as long as 
petals, those opposite petals united with them to %, with large anthers rounded 
at the peduncle apex; follicles ca. 4 mm long with straight hard beak; 
seeds oblong-lanceolate, brown, ca. 0.5 mm long. July— August. 

Mostly sandy or solonetz soils, pine forests; as a weed in fields, among 
shrubs in forests.— European part: Lad.-Ilm., U. Dnp., U. V., M. Dnp., 
V. -Don, L. Don, V. -Kama, Transv., Bl. Gen. distr. : E. Eur. (Poland, Finland). 
Described from Europe. Type in London. 

Note. The variety stepposum A. Bor. can be separated in the 
southeast of the European part of the USSR. Its characters are as follows : 
roots tuberculate, passing abruptly into slender roots; internodes scattered; 
leaves ovate, 2—3 cm long, 1—1.5 cm broad, amplexicaul, subentire or 
shallowly sinuate -dentate; inflorescence rather loose; pedicels longer 
than flowers; flowers small, pale yellow or greenish; sepals acute, 
x / 4 — a / 3 as long as petals; petals lanceolate, ca. 1 mm broad, 2—3 mm long. 

Forb -grass associations of steppe and forest -steppe zone, solonetz 
soils, river floodplains, meadows, among shrubs, forest clearings; as a 
weed in fields. — European part : V.-Don,, L. Don, V. -Kama (Sterlitamak), 
Transv.; W.Siberia: U. Tob. (Kustanai); Centr.Asia: Ar. -Casp. (Aktyubinsk). 
Owing to the presence of transitional forms it is impossible to isolate 
var. stepposum as a separate species. 

6. S. caucasicum (Grossh.) A. Bor. sp. nova in Addenda VIII, p. 366. — 
S. maximum var. caucasicum Grossh., Fl. Kavk. II (1930) 226. — 
S. telephium auct. s. lat. 

Perennial; roots thickened, fusiform, robust; stems 30— 70 cm high, 
ascending, strong, rounded, green or dark purple; leaves decussate, 
large, 5—7 cm broad, 3—5 cm long, broader than long, cochlear -concave, 
obovate, deeply cordate, amplexicaul, with large auricles, green or dark 
red, especially along the veins, obscurely and coarsely sinuate, obtuse; 
inflorescence corymbiform -paniculate, 5—12 cm broad, the lateral branches 
arising from axils of 2—3 upper leaf pairs; inflorescence branches long, 
many -flowered; pedicels as long as flowers, thick, strong; bracts fleshy, the 
lower larger, ovate -rounded, the upper small, oblong to lanceolate -terete; 
flowers 5-, sometimes 4 -merous, cup-shaped; calyx light green, with very 
short tube, fleshy, with triangular acute sepals, 1 /^ — % as long as petals; 
petals 0.5 cm long, greenish white, ovate, acute, with constricted carinate 



45 



58 



61 



apex, reflexed from the middle, the apex curving upward; stamens 10, with 
white filaments and yellow anthers, the stamens opposite to petals as long as or 
slightly shorter than them, those opposite to sepals slightly longer than petals; 
hypogynous scales yellow, obtuse, entire at the apex, twice as long as broad; 
follicles greenish white, with short reflexed beak, twice as long as broad, 
triquetrous, thickish, not closed, slightly shorter than petals; seeds small, 
ca.0.5 mm long, ovoid, subobtuse, brown. July — September. (Plate IV, 
Figure la). 

Dry stony and calcareous slopes in mixed deciduous forests, at 
1,000-2,500 m. Caucasus: Cisc, W. E., and S. Transc, Dag. Endemic. 
Possibly in Arm. -Kurd. Described from the Caucasus (Akhalbalaki). 
Type in Leningrad. 



Series 2. Viviparae A. Bor. Caudex short, with or without tuft of short 
roots or obsolete; leaves in whorls of 3—5, elongate-lanceolate; white, 
fleshy viviparous buds in axils of leaves and on rhizome. 

7. S.verticillatum L., Sp.pl. (1753) 430; Maxim, in Bull. Ac. Pet. XXK 
(1884)139; Berger in Engl. u. Pr., Nat. Pflanzfm. 18a (l 930) 444; Kom. et 
Alis., Opr. rast. Dal'nevost. kraya I, 596.— S.telephium ssp.verticil- 
latum Frod. in Acta Horti Gothoburg. V (1930) 64.- Ic: L., Amoen. 
acad.II (1787) 252, tab. 4, f. 14; Praeger in Journ. Hort. Soc. Lond. (1921 ), 
f. 44; Frod., 1. c, pi. XXV, textfig. 254-259. 

Perennial; caudex very short, woody or obsolete, with tuft of short, 
slender roots; stems 60—80 cm high, mostly solitary, erect, simple; 
leaves in whorls of 4 or 5, the lower partly in threes and opposite in pairs, 
the upper in whorls of 3—5, oblong -lanceolate or oval-oblong, distinctly 
petiolate, irregularly obtusely dentate, longer than internodes, sometimes 
with black dots, pale green, 5 — 8 cm long, 1.5—3 cm broad; viviparous 
white fleshy buds developing in leaf axils toward fall and growing during 
the fall; inflorescence compound, paniculate -corymbiform, dense, compact, 
4—8 cm in diameter; lower short inflorescence branches arising from 
leaf axils; bracts oval; flowers ca. 4— 5 mm long; sepals triangular, acute, 
V 4 — Y 3 as long as petals, shortly connate at base into a tube; petals oblong- 
lanceolate, acute, greenish white; stamens opposite to sepals slightly exceeding 
petals, the other 5 shorter, inserted almost at base of petals, with rounded 
reddish anthers and pale green filaments; hypogynous scales twice as long 
as broad, yellow, linear-cuneate; follicles oblong, about as long as petals, 
long-pediceled, with short beak; seeds 1.5 mm long, oblong, subobtuse. 
July — September. 

Riverbank thickets, river alluvium, meadows, valleys. — Far East: 
Kamch., more often in the South, solitary or in small groups; Sakh. 
(between Kami-Onori and Onori), west coast of Sakhalin, Revun River ravine. 
Gen. distr. : Jap. -Ch. : Hokkaido, Honshu; Sakhalin, Kurile Islands, Korea. 
Described from Siberia. Type in London. 

Note. S.verticillatum has been erroneously recorded by Linnaeus 
for S. Eur. He probably had in mind the cultivated species. 



46 




PLATE IV. 1-Sedum c a uc a si c u m A. Bor.: a) flower; 2 - S. pa llesce ns Freyn; 3 - S. p lu r i c a u le 
Kudo: a) flower; 4 - S. populi f oli urn Pall.: a) flower; 5 - S.tiyb ridum L: a) flower. 



47 



62 



8. S. viviparum Maxim, in Bull. Acad. Sc. Petersb. XXIX (1884) 137; 
Kom., Fl. Manch. II, 390; Kom. and Alis., Opr. rast. Dal'nevost. kraya I, 595. - 
S.telephium ssp.viviparum Frod. in Acta Horti Gothoburg. V (l 930 ) 
65.- Ic: Frod., 1. c, pi. XXVI, textfig. 260-268. 

Perennial; caudex very short or obsolete, with a tuft of short roots; 
fleshy white buds in axils of leaves and on rhizome; stems (17) 20—30 (35) cm 
high, mostly solitary, sometimes several, erect, simple; leaves in whorls 
of 3 or 4, some in pairs, dark green, lanceolate or ovate -oblong, somewhat 
rounded basally, with short cuneate petiole, very conspicuously veined, 
subobtuse, obtusely short -toothed from margin nearly to base, 3—4 (5) cm 
long, 1.2—1.5 cm broad, usually shorter than internodes, without black 
dots; inflorescence corymbiform -paniculate, compact, many -flowered, 
with very short pedicels and abortive flowers, bracteate; flowers small, 
open campanulate, ca. 3 mm long; sepals subobtuse, erect, ovate, x / 3 as 
long as corolla; petals oval or ovate, subobtuse, 3 times as long as 
broad, white or greenish, drying slightly pinkish; stamens opposite to petals 
about as long as or shorter than petals or shorter, the other 5 somewhat 
shorter than and inserted at base of petals, with subglobose pale 
anthers; hypogynous scales broader at the apex, thickened, curved; 
pistils shorter than stamens, ovate, with linear, reflexed style, half as long 
as ovary; seeds small, ovoid. August — September. 

Mixed forests, moist sites, rocks, sandy sites, always in the shade. 

Far East: Uss. (Vladivostok). Gen. distr. : Jap. -Ch. (N. Korea, Japan, 
N. Manchuria). Described from the Sedemi River, SE Manchuria. Type 
in Leningrad. 



Series 3. Fasciculatae A. Bor. Rhizome short, woody, with tuft of 
short roots; leaves alternate or opposite, flat, large, orbicular to elliptic. 

9. S.ussuriense Kom. in Bull. Jard. Bot. Petersb. XVI (1916) 170; 
Kom. and Alis., Opr. rast. Dal'nevost. kraya I, 596.— S.telephium ssp. 
purpureum var. orientale Frod. in Acta Hort. Gothoburg. V (1930) 66. — 
Ic: Kom. and Alis., 1. c, tab. 179. 

Perennial; caudex short, woody, multicipital, with a tuft of slender roots 
roots strong, terete or fusiform, but not thickened; all stems fruiting, 
ascending or erect, in groups, 20—60 cm long, leafy; leaves orbicular or 
broadly oval, amplexicaul, opposite, the lower alternate, sinuate -crenate, 
fleshy, subobtuse, 3—5 cm broad, 3—5 cm long; inflorescence dense, 
corymbiform-paniculate, with compact, thick branches; flowers mostly 4-, 
less often 5-merous, purple; calyx connate at base, with 4 oblong -ovate, acute 
lobes, 1 / i as long as corolla; petals oblong, ca. 3—4 mm long, acute, 1 -nerved; 
stamens 8—10, of which 4 (5) slightly exserted, about as long as petals, the 
4 (5) opposite to petals shorter, with purple anthers; pistils usually 4,- keeled on 
the outside, laterally inflated -convex, falcate, as long as or slightly longer 
than corolla; style short, erect or slightly curved. August — September. 
Fr. September. 

Seashore rocks. — Far East: Uss. (Peter the Great Bay, Suchan River 
valley). Endemic. Described from the vicinity of Vostok Bay. Type in 
Leningrad. 



48 



10. S.pallescens Freyn in Oest. Bot. Zeitschr. XLV (1895) 317; Kom. and 
Alis., Opr. Dal'nevost. kraya 1,596.— S.fabaria forma floribus lacteis 
Maxim., Fl. Amur. (1859) 115, p. p.— S.telephium var. pallescens 
Kom., Fl. Manshur. II (1904) 393.— S.telephium var. albif lorum Maxim. 
in Bull. Ac. Sc. XXIX (1884) 142.- S.albiflorum Maxim, in Kom. and 
Alis., 1. c. — S. t e 1 ephium ssp. p u r pu r eu m var. o r i e nt a 1 e Frod. in 
Acta Horti Gothoburg. V (l 930 ) 66.- S.leptorhizum Fisch. et Mey . 

in sched. - Ic. : Frod., 1. c, pi. XXIV. 

Perennial; roots slender, cordlike, arising in a tuft from a short, 
inconspicuous woody rhizome; stems densely leafy, erect, 30—60 cm high; 
leaves alternate, sometimes subopposite, elliptic -lanceolate, 3—7 cm long, 
0.7—2.5 cm broad, with cuneate base, sessile or petiolate, entire or 
irregularly sinuate -dentate, remotely and shallowly dentate in upper 
part of leaf, with numerous dark reddish dots (glands), pale green; 
inflorescence a corymbiform umbel, 3—5 (9) cm in diameter, 2.5 (7) cm 
long, arcuate, densely branching, compact; lower branches spreading, in 
axils of upper leaves; pedicels shorter ihan flowers; flowers 5 -merous, 
5— 6.5 mm long; sepals lance -linear, acute, J / 4 as long as petals; petals 
whitish or pinkish, erect, divided, lanceolate, acute; stamens 10, those opposite 
to petals shorter than and adnate to them to x / 3 , those opposite to sepals 
as long as or slightly longer than petals, lanceolate-elliptic, with filiform, 
reflexed long beak, green; seeds small, ca. 1 mm long, acute, brown. 
Fl. July — August, fr. August — September. (Plate IV, Figure 2 ). 

Wet and dry meadows, along rivers, riverbanks slopes, peaty riverbank 
meadows, sandy and sandy -gravelly soils.— E.Siberia: Dau.; Far East: 
Ze.-Bu. Described from Dauria. Type in Vienna. 

11. S. eupatorioides * Kom. and Alis., Opr. Dal'nevost. kraya I (1931 ) 
601. — S. t elephium var. e up a t o r io id e s Kom., Fl. Manch. II (1904) 
393.- S. fabar ia forma floribus lacteis Maxim., Fl. Amur. (1859) 115. — 
S. desmeti Palib. in sched. 

Perennial; roots slender, cordlike, arising in a tuft from a short 
woody rhizome; stem erect, densely leafy, 60—100 cm high, robust, strong; 
leaves alternate, subsessile, oblong-ovate, cuneate toward base, irregularly 
sinuate -dentate at the apex, (5)7—10 cm long, 1.5—4 cm broad, with abundant 
dark elongate -punctate glands below; inflorescence dense, 7—10 (l2)cm 
in diameter, 10—30 cm long, compound, umbellate -paniculate, many -branched, 
with up to 10 spreading branches, each bearing corymb; flowers 5 -merous, 
as long as or longer than pedicels; calyx Y 4 as long as corolla, with 
lanceolate -triangular, acute sepals; petals white or pink, 6—6.5 mm long, 
free, broadly lanceolate, acuminate, always erect; stamens 10, of which 
5 scarcely exserted from corolla, the 5 opposite to petals shorter than corolla; 
anthers yellow, later darkening, rounded; pistils oblong-lanceolate, with 
short, sometimes slightly curved style; fruit scarcely exserted from corolla, 
oblong-lanceolate, with short beak; seeds ca. 1 mm long, oblong-lanceolate, 
brown. July— August. 

Dry meadows and slopes, moist meadows, plowland. — Far East: Uss. 
(Amur River, near Khabarovsk, Vladivostok district, Suchan River basin, 
Lake Khanka). Gen. distr. : Jap.-Ch., Manchuria (Khanshakheza station); 
N. China, Korea. Described from the vicinity of Khabarovsk, from the 
banks of the Yalu River. Type in Leningrad. 

* This name was given on account of the large inflorescence resembling that of Eupatorium. 

49 



64 



Note. This species is closely related to S.pallescens Freyn, from 
which it is distinguished by its greater dimensions and its distribution 
area. 

12. S. alboroseum Baker in Saunders, "Refug. Bot." I (1868) tab. 33; 
Maxim, in Bull. Acad. Petersb. XXIX (l 884) 140 ; Kom., Fl. Manch. II, 391 . - 
S. telephium ssp. a lb o r o s e u m Frod. in Acta Horti Gothoburg. V (l 930) 
61.— Ic: Praeger in Journ. Hort. Soc. Lond. (l 921 ) f. 41 ; Regel, Gartenflora 
tab. 709, f. 4-5; Saunder, 1. c. 

Perennial; roots cordlike (according to Praeger — thickened), arising 
in a tuft from a short rhizome; stems 35— 60 cm high, hollow, yellowish, 
erect, robust; leaves opposite, sometimes in threes or approximate, 
alternate, ovate, with broadly cuneate base, subacute, largely obtuse, 
broadly dentate, (3)5— 10 cm long, 2— 4 cm broad; inflorescence dense, 
compact, sometimes with lateral branches, corymbiform -paniculate, 
2—6 cm in diameter; calyx 2 mm long, connate to the middle into a tube 
half as long as corolla, with broadly triangular sepals; petals 5, dry, 
whitish or pinkish, free, oblong-ovate, ca. 5 mm long, acute; stamens 10, those 
opposite petals shorter than and united with them to l /^—%, those opposite 
sepals as long as or slightly longer than petals; pistils lanceolate 
with erect, filiform style, gibbous on the back; seeds 1 mm long, oblong, 
brown. August — September. 

Stony and sandy soils at moistened sites on seashores and in river 
valleys.— Far East: Sakh. (Korsakovo; Tarantomari to the south of Mauko, 
Tym River valley). Gen. distr. : Jap.-Ch., Korea, Japan, Manchuria, China, 
Sakh. Described from Japan. 

Note. Specimens from Sakhalin Island and Korea differ somewhat from 
S. alboroseum (Maximowicz herbarium), mainly in leaf shape. It may 
be that the Sakhalin specimens should be referred to a new species, but 
lack of material does not permit this. The true S. alboroseum is 
cultivated as an ornamental plant. 



Subsection 2. HUMILICAULIA Praeger, 1. c, 96. - Stems weak, not erect, 
5—25 cm high, branching. 



Series 1. Repentes (Praeger) A. Bor.; Praeger, 1. c, 96, emend, pro gr. — 
Rhizome branching, cespitose; stems ascending, prostrate or creeping, 
branching. 

13. S. ewersii Ldb., Fl. alt. II (1830) 191; Fl. Ross. II, 182; Maxim., 
Bull. Ac. Petersb. XXIX (1884) 136; Kryl., Fl. Zap. Sib. VI, 1410; Fedtsch. 
O. et B., Consp. Fl. Turk. 3 (1909) 72.- S.altaicum Steph. in Sched. - 
S.azureum Royle III., Bot. Himal. I (1839) 223.'- Ic. : Ldb., Jc.pl. 
Fl. Ross. I (1829) tab. 48; Royle, 1. c, tab. 48, f. 2; Praeger in Journ. Hort. 
Soc. Lond. (1921) f. 45,46; Frod. in Acta Horti Gothoburg. V (1930) pi. 
XXI, textfig. 228-235. 

Perennial; rhizome branching, woody, cespitose, giving rise to numerous 
fruiting and sterile shoots; roots slender, cordlike; stems numerous, glabrous, 



50 



reddish woody at base and branching, spreading, ascending and rooting, 
10—20 (25) cm high; leaves opposite, glaucous -green, thickish, punctate, cordate 
at base, broadly ovate or suborbicular, bluntly short -acuminate, grayish, 
1—5 — 2 cm long and about as broad, obscurely denticulate, the lower broadly 
elliptic, often with brown spots; inflorescence corymbiform, compound, 
ca. 2—3 cmbroad; calyx with 5 lanceolate lobes, ca. 2 mm long; petals 5, twice as 
long as calyx, elliptic, lanceolate, 4—5 mm long, acute, pink or light purple; 
stamens 10, slightly longer than petals, with blackish anthers; hypogynous 
scales 0.5 mm long, oblong, slightly emarginate; follicles erect, 3—4 mm 
long, with short, reflexed beak; seeds small, ca.0.5 mm long, lanceolate, 
brown. July— August. 

Rocks, stony slopes, up to 3,500 m, reaching upper part of alpine 
zone.— W.Siberia: Alt.; Centr. Asia: Dzu. -Tarb. (Saur, Tarbagatai, 
Dzungarian Ala-Tau), T. Sh., Pam. -Al. Gen. distr. : Dzu.-Kash., W. Mong., 
Ind.-Him. Described from Altai. Type in Leningrad. 

14. S. pluricaule Kudo in Journ. Coll. Agric. Sapporo XII (1923) 40; 
Kom. and Alis., Opr. Dal'nevost. kraya I, 596.— S. cyaneum auct., non 
Rud. — S.telephium var. pluricaule Maxim, in Bull. Ac. Petersb. 
XXIX (1884) 142.- Ic: Miyabe and Miyake, Fl. Sach. (1915) pi. 6. 

Perennial; roots slender, cordlike; rhizome slender, cespitose, creeping; 
stems numerous, branching, with ascending branches, densely leafy, 5—7 cm 
long, rooting at the nodes; leaves opposite, some alternate, sessile, connate 
at base, somewhat tapering, broadly ovate, orbicular or obovate, entire, 
7—10 mm long, 5—8 mm broad, without red dots, obtuse, thickened at apex; 
inflorescence an umbellate corymb, many-flowered, ca. 2 — 3 cm in diameter, 
leafy; flowers ca. 5 mm long; pedicels shorter than or as long as flowers; 
calyx with 5 triangular, linear -subulate, acute sepals, 1 -nerved, 1.5—2 mm 
long, ca. 0.8 mm broad; petals 5, ca. 5 mm long, 2—2.5 mm broad, oblong- 
lanceolate, reclinate, acuminate, purple, 1 -nerved; stamens 10, with equal 
reddish filaments, filiform, the stamens slightly shorter than or as long 
as petals, with rounded, dark purple anthers, the 5 opposite to petals 
adnate to them to V 4 ; pistils 5, lanceolate, ca. 4 mm long, with slender, 
filiform, curved style; hypogynous scales small, trapezoid, 0.8 mm long, 
1 mm broad; follicles oblong -fusiform, many-seeded; seeds light brown, 
ovoid -oblong, small. July— August. (Plate IV, Figure 3a). 

In groups on eroded rocks, bald mountains, rock crevices, steep slopes, 
seashore cliffs, among Siberian dwarf -pines. — Far East: Sakh. : Pilvo- 
Porokotan, along the Engranvis River, Ikhdam, on mossy summits near 
the post of Due, Nabil Range, banks of the Nampi River, Ola District. 
Gen. distr. : Sakh. Described from Sakhalin. Type in Japan (?). 

15. S. cyaneum Rud. in Mem. Ac. Petersb. (1809) 341; Maxim, in Mel. 
Biol. XI (1873) 774; Ldb., Fl. Ross. II, 182; Maxim., Pr. Fl. Amur. (1859) 115; 
Kom. and Alis., Opr. rast. Dal'nevost. kraya I, 596; Berger in Engl. u. Pr., 
Nat. Pflzfm. 18a (1930) 445.- S.lilacinum Ldb. in Mem. Ac. Pe'tersb. V 
(1815)535; Chamisso in Linnaea VI (1831 ) 550. - S. d a hu r i c u m Steph. 

in sched. — S. hy p e r b o r eum Fisch. in sched. — Ic. : Rud., 1. c, tab. 1 1 ; 
Regel, Gartenflora tab. 972, f. 2; Praeger in Journ. Hort. Soc. (l92l) f. 53; 
Frod. in Acta Horti Gothoburg. V (192 9) pi. XXII, textfig. 236-41. 



51 



67 



Perennial; rhizome slender, creeping, prostrate, cespitose, with a row 
of slender roots; stems rooting in lower part, prostrate or creeping, 
ascending, branching, with ascending branches, reddish, with sterile and 
fruiting shoots, 5—7.5 (—12) cm high; leaves alternate, glaucous, with reddish 
dots, transparent at the margin, flat, fleshy, entire, sessile, oblong-lanceolate 
to sublinear, ca. 1 cm long, 2—3 cm broad, crowded on sterile shoots, 
subobtuse; inflorescence a corymbiform umbel, few-flowered, leafy; 
pedicels about as long as flowers; bracts of same shape as cauline leaves; 
sepals 5, narrow, oblong, acute, grayish-reddish, about half as long as 
petals, 3 mm long; corolla campanulate, pink -lilac or bluish; petals 5, 
oblong, acuminate, nodding, ca. 5— 6 mm long, 1.5—2 mm broad; stamens 10, 
with subulate dark purple filaments; 5 stamens adnate to petals to 1 / z , 
exceeding petals, the 5 opposite sepals longer; anthers rounded, 
angular, 4 -segmented, dark, with yellow pollen; hypogynous scales 5, small, 
oblong or cuneate-linear, entire; pistile 5, approximate, gibbous at 
base, reddish, with subulate, recurved style; follicles lanceolate, shorter 
than petals. Fl. June— August, fr. August— September. 

Stony soils, slopes, taluses, pebbles. — Arctic: An., E.Siberia: Lena-Kol. 
(Lake Dendyu, Batom River, an affluent of the Maya River); Far East: 
Okh., Kamch, Uda, Uss., Sakh. Gen.distr.: Sakh. (S.). Described from 
E.Siberia (Okh.). Type in Leningrad. 

*S. anacampseros L., indicated for the southern part of Podolia 
(Eichwald), the [former] Don Region* (Guldenstedt), and the Ukraine 
(Besser), does not in fact occur in the USSR; this is a S. European mountain 
plant, growing in the Pyrenees, the southern Alps, and Apennines. 

*S. fabaria Koch, has been erroneously indicated for the USSR; this 
species does not extend farther east than the Carpathians; the specimen 
recorded as S. f ab a r i a should, in fact, be classified as S. pu r pu r eum. 



Section 2. POPULISEDUM Berger in Engl. u. Pr., Nat. Pflzfm. 18a (1930) 
446.— Flowers white or pink, 5 -merous, bisexual; perennial, under shrubs, 
branching and woody from base, with woody roots; stems annual only in 
the upper, herbaceous part; leaves flat, petiolate, ovate, large, unequally and 
obtusely dentate. A single species standing in isolation from other Sedum 
species. 

16. S. populifolium Pall., Reise III (1776) Anhang, 730; Lin., fil. Suppl. 
(1781)242; Ldb., Fl. Ross. II, 180; Kryl., Fl. Zap. Sib. VI, 1408. - Ic. : Pallas, 
1. Ctab.O, f. 2; Praeger in Journ. Hort. Soc. (l 921 ) f. 78; Frod. in Acta 
Horti Gothoburg. V (1930) pi. XX, textfig. 211-21 9. 

Perennial; rhizome branching, cardlike, woody; stems branching, woody 
at base, spreading and partly prostrate, with brownish-gray bark, 15—35 cm 
high; leaves alternate, long -petioled, ovate or oblong -ovate, the lower 
cordate, with large, unequal obtuse teeth along the whole margin.the blade 
1.5—4 cm long, 1—2.5 cm broad, about twice as long as petioles, less often as long 
as petioles; inflorescence terminal, many -flowered, corymbiform -paniculate; 






[Part of Rostov Region since 1924.] 



52 



68 



pedicels as long as or longer than flowers; sepals 1.5—2 mm long, ovate- 
lanceolate, acute, connate at base; petals oblong-lanceolate, white or 
pinkish, 3—4 times as long as calyx, ca. 5—6 mm long and 2 mm broad; 
stamens as long as petals, with dark red anthers; hypogynous scales 
less than 1 mm long, 1.5 times as long as broad; follicles ca. 4 mm long; 
beak ca. 1 mm long, slightly reflexed; seeds small, ovoid, obtuse. 
June — August. (Plate IV, Figure 4a). 

Shady, moist sites, rocks and stony slopes, stream banks. W. Siberia: 
Alt. (E.), Ang. -Say. (Sayans). Gen. distr. : Tuva ASSR. Described from 
the Sayans. Type in London. 



Section 3. AIZOON Koch, Syn. (1837) 259. - Flowers yellow, bisexual, 
5-merous; follicles connate at base; rhizome short and thick, or else 
elongated, branching, and then leaves evergreen; roots not tuberous; 
leaves flat, alternate, oblong-oval to linear or lanceolate, dentate only in 
upper part of leaf, subcoriaceous. 

Species distributed in eastern Asia. 



Series 1. Glabrae A. Bor. — Glabrous plants. 

17. S.aizoon L. Sp.pl. (1753) 430; Ldb., Fl. Ross. II, 183; Maxim., Pr. 
Fl. Amur. (1859) 115; Turcz., Fl. baic.-dahur. 1,436; Maxim, in Bull. Ac. 
Petersb. XXIX (1884) 143; Kryl., Fl. Zap. Sib. VI, 1411 ; Kom. and Alis., Opr. 
rast. Dal'nevost. kraya I, 601 ; Kom., Fl. Manch. II, 395; Berger in Engl. u. 
Pr., Nat. Pflzfm. 18a (1930) 448.- S. sajanense Pall, ex Ldb., Fl. Ross., 
1. c.-Ic: DC, PI. Grass, tab. 101; Praeger in Journ. Hort. Soc. (l 921 ) 
f. 54a, 55, 56; Kom. and Alis., 1. c, 597, tab. 179; Frod. in Acta Horti 
Gothoburg. VI (l 931 ) pi. XLIII-XLV, textfig. 599-603. 

Perennial; rhizome short; stems 1—3,25— 45 cm high, fruiting and 
sterile, densely leafy, glabrous, erect, not branching; leaves alternate, 
elongate -lanceolate to linear, firm, subcoriaceous, unequally serrate - 
dentate, subacute, cuneate at base, 5—8 cm long; inflorescence a corymbiform 
umbel, sometimes paniculate, many-flowered, with horizontal branches, 
compact, flat, surrounded by leaves; sepals green, ovate at base, long- 
tapering in upper part, subulate, fleshy, / 4 — 1 / 3 as long as corolla, ca. 2 mm 
long, obtuse; petals golden-yellow, apiculate, narrow, elliptic -lanceolate 
(6)7—10 mm long; stamens 10, all equal, shorter than petals, inserted 
somewhat above base of petals; ovaries connate at base, ovate, convex 
ventrally, with subulate style; hypogynous scales green, barely discernible, 
subquadrate; follicles many -seeded, stellately spreading, laterally 
compressed, ca. 7 mm long, with short straight beak; seeds brown, ellipsoid, 
ca. 7 mm long, with short straight beak; seeds brown, ellipsoid, less than 
1 mm long. Fl. from middle of June to late fall. 

Dry grassy slopes, dry shrub thickets, dry meadows, rock streams, 
sandy cliffs.— W.Siberia: Ob (city of Tara), Irt. (widely distributed); 
E.Siberia: Yenis. (many), Ang. -Say., Lena-Kol. (rarely to Nizhne-Kolymsk), 
Dau. (many); Far East: Okh., Kamch., Ze.-Bu., Uss., Uda, Sakh. Gen. distr.: 
Jap.-Ch., Mong. (Tuva ASSR). Described from Siberia. Type in London. 



53 



69 



Note. Extremely polymorphic and undoubtedly a collective species. 
The leaves are especially variable. Several forms can be distinguished: 
the narrow -leaved f. angusti folia A. Bor., occurring in the western 
part of the distribution area (Ob, Irt., Yenis., Dau.), and the broadleaved 
f. latifolium A. Bor., growing in the eastern part (Uss., Ze.-Bu., Korea, 
Sakh., Jap.). A mostly branching form with elliptic, coarsely unequally 
uncinate -dentate leaves is confined to the Maritime Territory. The 
presence of a series of transitional forms renders the separation of new 
species more difficult. Cultivated plants. 

18. S. hyperaizoon Kom. in Bull. Jard. Bot. Princ. XXX, f. 1-2 (1931 ) 
201. - S. aisoon var. latifolium Maxim., Prim., Fl. Amur. (1859) 115; 
Bull. Ac. Pet. XXIX (1884) 143. 

Perennial; rhizome short, multicipital; stems solitary, erect, robust, 
ca. 6 mm thick, to 85 cm high, not branching, glabrous; leaves alternate, 
ovate-lanceolate, 8—10 cm long, 3—4 cm broad, cuneate at base, short - 
petioled, coarsely serrate -dentate, obtuse, firm, subcoriaceous; inflorescence 
obconical, broad, corymbiform -umbellate, compact, with short thick 
branches, spreading subhorizontally, surrounded by numerous bracts; sepals 
green, ovate or triangular -acuminate, attenuate at the apex; petals narrowly 
lanceolate, ca. 6 mm long, 3—4 times as long as calyx; stamens all equal, 
shorter than petals; follicles stellately spreading, gibbous ventrally, 
laterally compressed, with erect subulate beak Y 3 as long as follicle; 
seeds small., less than 1 mm long, ellipsoid, brown. June. 

Rocks.— Far East: Uss. (on the shore of Russkii Island on rocks 
among oak groves; Shufan on the Sungacha River). Endemic. Described 
from the Far East (Russkii Island). Type in Leningrad. 

Note. S. hyperaizoon Kom. is closely related to the polymorphic 
S. a i z o o n L. It is an extreme form of the broadleaved S. a i z o o n L. ; 
very closely related to S. maximoviczii Rgl., in cultivation in Japan, 
and overwintering well in Leningrad. 

19. S. kamtczaticum Fisch. in Ind. Sem. H. Petropol. 7 (1840) 54; 
Maxim, in Bull. Acad. Pet. XXIX (1884) 145; Ldb., Fl. Ross. II, 182; Kom., 
Fl. Manch. II, 398; Kom. and Alls., Opr. rast. Dal'nevost. kraya I, 601 ; 
Berger in Engl. u. Pr., Nat. Pflzfm. 18a (1930) 449.- Ic: Praeger in 
Journ. Hort. Soc. (l 921 ) f. 54c, 62. 

Perennial; roots cordlike, slender; rhizome strong, woody, elongated, 
with underground shoots; stems few, 30—40 cm high, erect or ascending, 
sometimes rooting in lower part, with evenly spaced leaves; leaves 
alternate, oblong-lanceolate, spatulate, with cuneate base, obtuse, coarsely 
and obtusely dentate along the margin only at the apex, entire at base, 
lower leaves almost all entire, dark green; inflorescence broad, loose, 
flatfish, corymbiform, with few-flowered short branches, 1—5 -flowered, 
surrounded by bracts surpassing the inflorescence; flowers sessile or 
short -pediceled; sepals ovate, with elongated apex; corolla stellate, 
orange -yellow; petals twice as long as sepals, lanceolate, apiculate; 
stemans as long as or somewhat shorter than petals, with orange anthers; 
pistils reddening after anthesis, connate to more than V 3 from base, shorter 



54 



than corolla; follicles united to more than 1 / 3 , bright red at maturity, 
stellately and horizontally spreading; seeds obovoid, dingy brown -yellow. 
June — September. 

Stony slopes.— Far East: Uss., Kamch., Sakh. Gen. distr. : Jap. -Ch. 
Described from specimen under cultivation. Type in Leningrad. 

Note. Often cultivated as an ornamental plant. Cultivated 
f. variegatum has white leaf spots. 

There are the following hybrids : l)S.aizoonX S.kamtczaticum, 
with leaves as in S.aizoon, inflorescence and flowers as in 
S. kamtczaticum; blooming like S.kamtczaticum 14 days earlier 
than S.aizoon (Praeger, 1. c, f. 57); 2) S.kamtczaticum X S. maxi- 
mowiczii Praeger. 

20. S.middendorfianum Maxim., Prim. Flor. Amur. (1859) 116; Ej. in 
Bull. Ac. Peters. XXIX (1884) 146; Kom., Fl. Manch. II, 397; Berger in 
Engl.u. Pr., Nat. Pflzfm. 18a (1930) 448.- S.aizoon var. middendor- 
fianum Frod. in Acta Horti Gothoburg. VI (l 931 ) 80. — Ic. : Praeger in 
Journ. Hort.Soc. (1921) f. 54, 59, Frod., I.e., pl.XLVIII, 1, textfig. 609-613. 

Perennial; rhizome creeping, woody, branching, long; stems numerous, 
fruticose, old stems persistent, straight, suberect, branching at base, 
glabrous, slender, leafy, 10—15 (30)cm high, with sterile, shorter shoots; 
leaves narrow to linear, the lower spatulate, other leaves linear -spatulate, 
with few obtuse teeth at the apex or obtusely crenate to the middle, 
canaliculate, subobtuse; inflorescence rather loose, paniculate-corymbiform, 
many -flowered, often with elongated -spreading, many -flowered ascending 
branches with scattered flowers; sepals 5, linear, subobtuse; corolla 
stellate, to 18 mm in diameter; petals lanceolate to linear -lanceolate, 
acuminate, twice as long as calyx, yellow, 5— 6 mm long; stamens 10, half 
as long as petals, with yellow filaments and orange anthers, those opposite 
to petals adnate to them to x / 4 of length from base; pistils adnate at 
base to slender styles; hypogynous scales small, very short, whitish, 
compressed, entire; follicles greenish, stellately spreading, 4 mm long, 
with very short beak, laterally compressed, lanceolate; seeds ovoid, small. 
June — August. 

Rock crevices, forests, stony soils. — E. Siberia: Lena-Kol., Dau.; 
Far East: Okh., Uda. Gen. distr. : Jap. -Ch., N. Manchuria. Described 
from Khadzhi (Sovetskaya Gavan). Type in Leningrad. 

21. S.hybridum L., Sp.pl. (1753) 431; Ldb., Fl. Ross. II, 183; Maxim, 
in Bull. Ac. Pet. XXIX (1884) 147; Kryl., Fl. Zap. Sib. VI, 1412; Turcz., Fl. 
baic.-dahur. 1,436; Berger in Engl.u. Pr., Nat. Pflzfm. 18a (1930) 449.- 
Anacampseros hybrida Flaw., Syn. ed. germ, (l 81 9) 123. — Ic. : 
Rchb., Ic. Fl. Germ. XXIII, tab. 64; Praeger in Journ. Hort. Soc. (l 921 ) f. 54e, 
65; Frod. in Acta Horti Gothoburg. VI (1931 ) pi. XLIX, textfig. 622-626. 

Perennial; rhizome long, branching, woody, cordlike, creeping, with 
slender roots; stems evergreen, not dying off, 15—20 (30) cm long, creeping, 
prostrate and rooting, profusely branching; sterile shoots densely leafy, 
short; fruiting shoots to 30 cm high, ascending, with smaller, remote 
leaves; leaves alternate, 1.5—3 (5) cm long, (0.8) 1— 2 cm broad, spatulate- 
elliptic, obtusely dentate on the margin, often with reddish teeth, gradually 



55 



tapering to a Long, cuneate, entire, petiole about as Long as the blade, 
subobtuse, glabrous, Fleshy; inflorescence ;i terminal dense paniculate - 
corymbiform cyme with elongated branches; sepals Lanceolate, subobtuse, 
pale green, ca. 3 mm Long, even, the a pox not elongated, connate at base; 
corolla yellow; petals twice as long as calyx, elliptic -lanceolate, acute, 
,,. recurved, ca. 6 mm long, 1.5 mm broad; stamens opposite to petals slightly 
shorter than them, those opposite to sepals about as long as petals, with yellow 
filaments and orange authors; pistils shorter than petals, lanceolate, 
with filiform style, pale green; bypogynous scales very short, shorter 
than broad, bluntly truncate at the apex; follicles elliptic, 7— -8 mm long, 
convex venl rally, obliquely declinate, stellately spreading, united to l / 3 
from base, green, reddish at the apex; seeds numerous, elliptic, obtuse, 
Less than I mm Long. Fl.from June, fr. July— August. (Plate IV, 
Figure 5a). 

Stony and gravelly soils, rock crevices, less often sandy ami pebbly 
bluffs; mainly mountain steppes, less often in southern parts o( the 
foresl .one. Kuropenn part: Urals (central and southern parts of the 
mountain range); W.Siberia: Irt., Alt.; E. Siberia: Yenis., Ang. -Say., 
Lena-Kol.j Centr.Asia: Ar.-Casp. (Akmolinsk), T.Sh. Gen.distr.: 
\. Mong., Tuva. Described from S.Urals (Tataria, ad radices montium 
uralensium). Type in London. 

22, S. litorale Kom. in Bull. Jard. Bot. Princ. XXX, f. 1 -2(1931)201. 
Perennial: rhizome elongated, creeping, simple, with a monocephalous 

root; stems st rnong, slightly flexuous, 32 cm long, lustrous, glabrous ; 
Leaves in whorls of 3, the upper early deciduous, sessile, ovate or obovate- 
Lanceolate, 4 6 cm long, 1.8—3.5 cm broad, subobtuse, rounded at the apex, 
broadly serrate -dentate by uncinate adjacent teeth, with small white dots 
Oil the blade; ini ernodes on. I' cm Long, inflorescence compact, with 
angular branches; flowers short -pediceled or sessile, golden; sepals 
thickened, whitish, triangular; petals lanceolate, acute, ca. 4 mm long: 
follicles stellately spreading, rounded dorsally, keeled ventrally, with 
long horizontally declinate beak. August. 

Collected on sandy seashores.— Far East: Uss. (Popov Island in Peter 
the Greal Hay near Vladivostok). Endemic. Described from the Fat- 
East. (Popov Island). Type in Leningrad. 



Series 2. Villosae A.Bor.— Entire plant densely pubescent. 

23. S. selskianum Rgl. et l\laak in Rgl., Tent. Fl. Ussur. (1861) 66; 
Maxim, in Bull. Acad. Petersb. XXIX (1884) 145; Kom. and Alis., Opr. 
rast, Dal'nevost. kraya 1. 601 : Kom., Fl. Manch. 396; Berger in Engl. u. 
Pr., Nat. Pfl.-fm. 1 8a (1930) 449. - S. aizoon ssp. selskianum Frod. 
in Acta Horn Gothoburg. VI (1931 ) 80.- lc: Rgl., 1. c, tab. VI, f. 9-10; Mem. 
Ac. Sc. VII, tab. IV, 4 (l86l)j Rgl., Gartenflora (l 862) 169, tab. 361 ; 
Praeger in Journ. Won. Soc. (1921 ), f. 58; Frod.. 1. c, pi. XLVIII, 2; 
textfig. 614—621, 

Perennial; roots cordlike, numerous; rhizome short, woody; entire 
plan! covered with dense grayish pubescence; stems numerous reddish, 



56 



woody, erect or ascending, simple or branching, 35—40 cm high, leafy; 
leaves alternate, sessile, declinate, crenate -serrate, spatulately narrow- 
lanceolate or linear, subacute or subobtuse, entire and cuneate from middle 
to base, 3— 6 cm long, 0.3— 1 cm broad; Inflorescence corymbiform with 
many-flowered branches surpassing bracts, 3— 7 cm In diameter, leafy j 
flowers small, bright yellow, with very short pubesceni pedicels; sepals 
green, fleshy, usually glabrous or sparsely pubescent, trlangula r -lanceolate, 
subobtuse, erect, free to base, V 2 — 2 / 3 as long as corolla; petals ca. 5 nam 
long, yellow, broadly lanceolate, acuminate or apiculate; stamens all equal, 
not exserled, with yellow anthers; Collides small, rounded dorsal ly, dee linal.e 
at an obtuse angle from the middle, with long filiform beak; seeds light 
brown, less than 1 mm long, ovoid, obtuse. July —August. 

Dry rocky and stony slopes, deciduous forests and fields.— Far East: 
Uss. Gen.distr.: Jap. -Ch., Manchuria. Described from the Ussuri River. 
Type in Leningrad. 



Section 4. EUSEDUM Roiss., Fl. Or. II (1872) 775. - G e n u i n a Koch, 
Syn. Deutsch. u. Schweiz. fl. (1837) 259.— Plants with creeping rhizomes, 
mostly evergreen, cespitose at base; stems erect, creeping or ascending 
from base; flowering stems with sterile shoots; leaves mostly semiterete 
or scalelike and thickened, otherwise flat and spatulate but then flowers 
white or pinkish. Flowers white, rv.i\, pink, yellow. 



Subsection 1. SPATHULATA A. Bor.- Leaves opposite, flat, spatulate, 
oval or obovate; flowers white or pink. 



Series 1. Involucratae Maxim, ex Berger, 1. c, 449. — Plants without 
white underground buds. 

24. S. stevenianum Rouy et Camus, PI. Fr. VII (l 901 ) 94; Grossh., Fl. 
Cauc. II, 22G; Berger in Engl. u. Pr., Nat. Pflzfm. 18a (1930) 450; !«'rod. in 
Acta Hort. Gothoburg. VII (1932) 12.- S. r o s e u m Steven (non Scop. ) in 
Mem.Soc.Nat.Mosc.nl (1812)263; Ldb., Fl. Ross. II, 184; Boiss., Fl. Or. II, 
402.- Ic: Praeger in Journ. Hort. Soc. (1921 ) f. 113; Grossh. in Monit. 
Jard. Bot. Tiflis III, IV (1917) 135, f. 6. 

Perennial; roots fibrous; rhizome slender, branching, cordlike, cespitose; 
stems slender, profusely branching, weak, flexuous, appressed to the ground 
or ascending, creeping, glabrous, (2.5)3—10 cm long, fruiting and sterile; 
leaves opposite, more approximate on sterile shoots, flat, fleshy, sessile, 
glabrous, entire, ovate or spatulate, cuneate at base, obtuse, 2—6 mm long, 
1— 4mm broad; inflorescence umbellate, (l ) 3— 9 -flowered, 1—2 cm in 
diameter; pedicels about as long as calyx; sepals glabrous, connate at base, 
linear -lanceolate, obtuse, Y 3 — l / 2 as long as petals, green; petals pink 
or whitish, with pink keel, oblong-lanceolate, entire, subobtuse or with a 
mucro 5— 7 mm long; stamens 10, Y 3 as long as petals, with green filaments 
and pink or reddish anthers; hypogynous scales pale orange, suborbicular, 
emarginate, forming a pocket, with differentiated lateral walls; follicles 
green, erect, shorter than stamens; seeds small, ca. 0.7 mm long, brown, 
oblong-ovoid. June — August. 



57 



Rocky sites in the alpine zone.— Caucasus: Dag., E. Transc, possibly 
in Cisc. on the northern slope of the Main Range. Gen. distr. : Bal. -As. Min. 
(?). Described from the E. Caucasus, near the village of Budukh. Type 
in Leningrad. 

25. S. stoloniferum S.G. Gmel., Reise III (1774) 311; Ldb., Fl. Ross. II, 
185; Boiss., Fl. Or. II, 779; Hamet in Acta Horti Tiflis. VIII, 3 (1908) 8; 
Berger in Engl. u. Pr., Nat. Pflzfm. 18a (1930) 450.- S.ibericum Stev. 
in M.B.; Fl. taur. -cauc. Ill (1819) 312.- S.hybridum Urv. ex Boiss., 
Fl. Or. 11,779, non L. - Ic: Gmelin, 1. c, III, tab. XXXV, f. 2 ; Praeger 

in Journ. Hort.Soc. (1921) f . 1 1 1 ; Grossh. in Monit. Jard. Bot. Tiflis 
III-IV (1917) 135, f. 9; Frod.Acta Horti Gothoburg. VII (1932) pl.XXVHI, 
textfig. 369-375.- Exs.: Herb. Fl. Cauc, No. 324. 

Perennial; roots slender, fibrous; rhizome long, creeping, cordlike; 
flowering, erect, 17—35 cm long, glabrous or scabridulous along the ribs, 
sterile, ascending or prostrate, rooting, 5 — 12 cm long; leaves opposite, re 
remote, flat, with petioles 3—7.5 mm long, the blade 8— 15 mm long, 8— 12 mm 
broad, rounded -oval, rhomboid -spatulate or oval, with cuneate base, obtuse, 
broadly sinuate-dentate, with a very narrow, transparent, light stripe; 
young leaves with white dots, glabrous or pubescent; inflorescence 
umbellate, with divaricate, elongated, flexuous branches, leafy, with 
scattered flowers; bracts gradually tapering upward; flowers subsessile 
or short -pediceled, 0.2—0.5 cm long; sepals linear, subobtuse, Y 3 — x / 2 
as long as corolla, 2—3 mm long, green, connate at base; petals pink, 
linear-lanceolate, acute, 5— 8 mm long, 1.5 mm broad; stamens 10, about 
half as long as petals, the 5 opposite to the petals adnate to 7 3 of their 
length; anthers bright red, filaments pink; pistils pink, later turning 
white, convex ventrally, with short filiform style; hypogynous scales 
very small, subquadrate, slightly emarginate, pink; follicles many-seeded, 
„. stellately spreading, with 2 ventral protuberances; beak filiform; seeds 
small, ovoid, subobtuse, 0.5—0.9 mm long, 0.4 mm broad, tapering at one 
end, blackish-brown. June — August. (Plate V, Figure 8a, b). 

Typical forest plants, growing in the lower and middle mountain 
zones on stony soils, sometimes in subalpine meadows. Rarely noted 
as a weed of tea plantations and on mountain paths.— Caucasus: Cisc, 
W. and E. Transc, Tal., less in S. Transc, Dag. Gen. distr.: Iran.: 
Asterabad [Gurgan], Gilan, Mazanderan; Arm. -Kurd. Described from the 
province of Gilan in Iran. Type in Berlin. 

Note. In the Caucasus this species is associated with large forest 
tracts confined to the region of two relict forest floras: the Ponthian and 
the Hyrcanian (Grossheim). 

Economic importance. Ornamental plants. 

26. S. spurium M. B., Fl. taur. -cauc I (1808) 352; Ldb., Fl. Ross. II, 183; 
Boiss., Fl. Or. II, 778; Shmal'g., Fl. I, 361 ; Grossh., Fl. Cauc II, 226; Hamet 
in Acta Horti Tiflis. VIII, 3 (l 308) 11; Berger in Engl. u. Pr., Nat. Pflzfm. 
18a (1930) 450.- Crassula crenata Desf., Choix Cor. Tourn. (1808) 76, 
tab. 58.- Sedum crenatum Boiss., Fl. Or. II (1872) 779. - S. d e nt at u m 
DC. Prodr. EI (1828) 403.- S. conge stum C.Koch ex Boiss., 1. c, 779.- 



58 



(75) 




PLATE V. 1-Sedum pe nt ape t a lu m A.Bor.: a) fruit; 2 — S.hispanicum L.: fruit; 3- 

s. pallidum M.B.: a) fruit; 4-S.rubrum (L.) Thell., in fruits: a) flower; 5 — S. tetra merum 

Trautv., in fruits: a) flower; 6 - S.lenkoran icum Grossh.:a) flower, b) pari oi Stem with Li ifl 

7 — S. subulatum (C.AM.) Boiss.: a) flower; 8 — S. stoloni fcr u in S.G„ Gmel.: a) Flower, I'" 1 fruit, 



59 



77 



S. la zi cum Boiss. et Huet, Diagn. Ser. 1, 2 (1856) 63.- S. ciliar e Sweet. 
Hort. Brit. (1827) 179. — A na camp s e r a s ciliar is Haw., Syn. pi. 
Succul. (1819) 12 9.- Ic: Praeger in Journ. Hort. Soc. (1921) f. 110; Rchb., 
Ic. Fl. Germ. 23 (l 898-1 899), tab. 46; Sims, Bot. Mag. (l 823) tab. 2370; Grossh. 
in Monit. Jard. Bot. Tiflis III - IV (1917) 135, f. 7.- Exs. : HFR. no. 1765. 

Perennial; rhizome creeping, long; roots slender, branching, fibrous; 
stems prostrate or ascending, the sterile shorter than the fruiting, 3—6 cm 
long, with more crowded leaves; fruiting stems 6— 20(25)cm long, puberulous 
or scabrous, with traces of fallen leaves; leaves fleshy, dark green, opposite, 
obovate-cuneate, 1—2.5 cm long, 0.5 — 1 cm broad, subobtuse, subobtusely 
dentate or crenate in upper part of leaves, puberulous, narrowly membranous 
and ciliate on the margin; inflorescence divaricate, umbellate-corymbiform. 
dense, mostly consisting of 4 strong, flexuous branches, with flowers in 
the bifurcations; flowers with bracts surpassing inflorescence, subsessile 
or short -pediceled, pink or purple, 10 — 15 mm long; sepals narrowly 
lanceolate, erect, green or reddish, fleshy, persistent in fruit, parted almost to 
base; corolla 2—2.5 times as long as sepals; petals lanceolate, acute, 
keeled, constricted at the apex, usually erect, entire; stamens 10, slightly 
shorter than petals, with red filaments and orange-red anthers, the stamens 
opposite the petals united with them to 1 / 3 of length from base; pistils 5, 
oblong, with erect style, pink; hypogynous scales whitish, broader than long; 
erect follicles not stellately spreading, reddish, oblong-ovoid, striped, 
1 mm long, 0.5 mm broad; seeds numerous, small, less than 1 mm long, 
subobtuse, oblong. June — August. 

Rocky sites, in the middle and upper mountain zones, in subalpine 
meadows.— Caucasus: mainly Cisc. and W. Transc, less in Dag. and 
E. Transc. Wild in M. Dnp. near Korostyshev and in Bl. near Uman. 
Gen. distr. : Arm. -Kurd. (Turkish Armenia). Described from the Caucasus. 
Type in Leningrad. 

Economic importance. The most widespread species under cultivation, 
grown as ornamental plants. Sometimes found wild. 

27. S.oppositifolium Sims, Bot. Mag. tab. 1807 (1815), non Hamet; 
Ldb., Fl. Ross. II, 184; Boiss., Fl. Or. II, 778; Grossh., Fl. Cauc. II, 227.- 
S. spur ium var. alba Trautv. in A. H. P. IV (1876) 370.- S. spur ium 
Hamet in Acta Horti Tiflis. VIII, 3 (1908) 11 (non M. B.). - S.spurium 
Frod.in Acta Horti Gothoburg. VII (1932) 10.- Ic. : Bot. Mag., I.e.; Rchb., 
Ic. Fl. Germ. XXIII (1898-1899) tab. 63; Grossh. in Monit. Jard. Bot. Tiflis 
III— IV (1917) 135, f. 7.- Exs.: Herb. Fl. Cauc, No. 321. 

Perennial; rhizome creeping, long; roots slender, fibrous; stems 
ascending, creeping at base, with traces of fallen leaves, pubescent, the 
fruiting stems 6 — 15 cm long, the sterile shorter; leaves opposite, obovate- 
cuneate, 1—2.5 cm long, 0.5 — 1.5 cm broad, short -petioled, subobtusely 



60 



truncate or subobtuse, coarsely crenate -dentate from the upper part to 
the middle, puberulous or glabrous, ciliate and narrowly membranous on 
the margin; inflorescence dense, umbellate, corymbiform, spreading, the 
bracts not surpassing inflorescence; flowers short -pediceled or subsessile; 
sepals lanceolate, obtuse, erect, light green; corolla 2—3 times as long 
as sepals; petals linear -lanceolate, acuminate, 0.7— 1 .3 mm long, white or 
pale yellowish cream-colored; stamens 10, with whitish filaments and 
yellow, later dark, anthers, slightly or markedly shorter than petals 
(ca. 2 / 3 as long); hypogynous scales emarginate, broader than long, attached 
to carpels by their base and lateral edges, forming a kind of pocket; follicles 
not stellately spreading, erect, with persistent calyx, green; seeds ovoid, 
less than 1 mm long, striped. July — August. Dry grassy, woodless, mainly 
southern and pebbly slopes and taluses; vertical distribution range large.— 
Caucasus: mainly E. and S. Transc, Dag., less in Cisc. and W. Transc, 
Tal. Gen. distr. : N.Iran. Described from the Caucasus. Type in London. 

Note. This species is very closely related to S. s p u r ium M. B., 
from which it is distinguished only by its white or pale yellow corolla, 
somewhat different leaf shape, and distribution area. S. oppositifolium 
Sims, occupies regions in the Caucasus where S. spurium does not occur. 
S. oppositifolium is a more xerophytic species, growing in drier 
regions in the E. and the S. Caucasus, while S. spurium grows mostly in 
the more moist N. and (partly) W. parts. In Daghestan and in the former 
District of Artvin, where both species occur, transitional forms with 
indeterminate coloration may be found (according to Grossheim). 

Economic importance. Widely cultivated, like S. spur ium, as an 
ornamental plant. 

28. S. involucratum M. B., Fl. taur.-cauc. I (1808) 352; Ldb., Fl. Ross. II, 
183; Boiss., Fl. Or. II, 778; Grossh., Fl. Cauc. II, 226; Hamet in Acta Horti 
Tiflis. VIII, 3 (1908)15; Berger in Engl. u. Pr., Nat. Pflzfm. 18a (1930) 
449. — S. s p u r iu m var. involucratum Frod. in Acta Horti Gothoburg. 
VII (1932) 11.- Ic: Gross, in Monit. Jard. Bot. Tiflis, III-IV (1917) 135, 
f. 7. 

Perennial; rhizome creeping, long; roots slender, fibrous; stems 
ascending, strong, simple, puberulous, the sterile ones 7—12 cm and the 
fruiting stems 10— 15 cm high; leaves opposite, obovate-cuneate, the blade 
1.5—1.8 cm long, 1.4—1.6 cm broad, with 4— 6 mm petioles, ciliate-margined, 
obscurely broadly sinuate, in upper part, truncate -obtuse, cuneate at 
base; inflorescence umbellate -corymbiform, dense, closely surrounded 
by large terminal leaves long-ciliate on their margin and often surpassingthe 
inflorescence; flowers ca. 10 mm long, sessile or with very short pedicels; 
sepals narrowly lanceolate, light green, 2 / 3 as long as petals, acuminate; 
petals narrowly lanceolate, acuminate, white or yellowish, uneven-margined, 
with very narrow white stripe; stamens 10, shorter than petals, more than 
half as long as petals, with white filaments and later -darkening anthers; 
hypogynous scales semiorbicular, deeply emarginate, broader than long; 
mature fruit erect, not stellately spreading, without ventral thickenings, 
their beak erect, linear; seeds ovoid, acute unilaterally, small, ca. 0.5 mm 
long, brown. July — August. 



61 



79 



80 






Stony soils in subalpine meadows up to 2,600 — 3,000111.— Caucasus: 
mainly Cisc. — northern slope of Alain Range, less in E. Transc. (Mt. Kazbek, 
Kobi), and Dag. Endemic. Described from the Caucasus (Kobi). Type 
in Leningrad. 



Series 2. Proponticae Berger., 1. c.,450.— Plants with spherical, white, 
underground buds. 

29. S. obtusifolium C. A. M., Enum. pi. Cauc. (l83l) 150; Ldb., Fl. Ross. II, 
184; Boiss., Fl. Or. H, 777; Hamet in Acta Horti Tiflis. VIII, 3 (1908), 1 1 ; 
Berger in Engl. u. Pr., Nat. Pflzfm. 18a (1930) 450.— S.gemmiferum 
Woron. in Izv. Kavk. Muz. V (1907) 205.— S. proponticum Aznavour in 
Bull. Soc. Bot. Fr. 44 (1897) 169. - Ic. : Frod. in Acta Horti Gothoburg. 
VII U932) pi. XXIX, textfig., 376-383; Grossh. in Monit. Jard. Bot. Tiflis. 
36 (1915) tab. 1; Grossh., 1. c, III, IV (1917) 135, f. 2; Praeger in Journ. 
Hort.Soc. (1921) f. 112.- Exs. : Herb. Norm. ed. Dorfler, No. 4871. 

Perennial; roots slender, fibrous; rhizome short, with spherical, light - 
colored buds, 5— 8 mm in diameter, later becoming detached and germinating; 
stems erect, villous, 10—25 cm high, strong; leaves opposite, flat, sessile, 
6—30 mm long, 5—18 mm broad, the upper oblong -oval, the lower ovate- 
rounded, larger, somewhat tapering toward base, obtuse, glabrous, reticulately 
red veins, denticulate, with a very narrow, light, transparent stripe; 
inflorescence umbellate, consisting of 2 or 3 obliquely ascending, leafy 
branches; flowers subsessile, with lanceolate bracts; calyx with oblong - 
lanceolate sepals, V 3 — 1 / 2 as long as sepals, glabrous; petals pink, lanceolate, 
entire, acute, 7— 8 mm long; stamens 10, 2 / 3 as long as petals; pistils 
oblong, as long as stamens, with short styles; hypogynous scales broader 
than long, slightly emarginate and tapering; follicles and basal protuberance, 
stellately spreading, lanceolate, with erect, subulate beak; seeds rounded - 
ovo id . May — Augu st . 

Mountain-xerophyte zone.— Caucasus: E. Transc. (Azerbajan), Tal. 
(Germin post, villate of Perimbal'). Gen. distr.: Arm. -Kurd.— possibly in 
adjacent dry regions of Iran, Bal.-As. Min. (Asia Minor, opposite 
Constantinople [Istanbul]). Described from Talysh. Type in Leningrad. 

Note. A number of authors — among them Meyer, who described 
S. obtusifolium, later Boissier (1872), Hamet (1908), Ledebour 
(1844—1846) — give an incorrect description of this plant. The error 
consisted in overlooking a character as important as the very characteristic 
villous pubescence. In addition, Boissier mistakenly records the corolla 
as being white instead of pink. Grossheim corrected these errors in 
the Flora of the Caucasus. 

30. S.listoniae Vis. in Inst. Sc. Venet. Mart. I (l84l) 21; Boiss., Fl. 

Or. II, 779; Berger in Engi. u. Pr., Nat. Pflzfm. 18a (1930) 450.— S.cariense 
Jaub. et Spach, 111. pi. Or. I (1842—1843) 16.- S.anatolicum C.Koch in 
Linnaea XIX (1847) 41.— S. obtusifolium var. listoniae Frod. in 
Acta Horti Gothoburg. VII (1932) 51.— Ic: Visiani, 1. c, tab. 6; Jaub. et Sp., 
I.e. tab. 7; Frod. pi. XXLX, textfig. 384-393. 



62 



Perennial; rhizome with small spherical buds and slender roots; 
flowering stems 10— 25 cm long, sterile stems 9— 12 mm long. Cespitose 
plants, ascending, glandular -puberulous; leaves opposite on the sterile and 
approximate on the fruiting stems, the lower leaves opposite, the upper 
remote, alternate, all flat, sessile, ciliate -margined and entire, spatulately 
obovate or oblong, obtuse, tapering toward base, 1.5—4 cm long, 1—3 cm 
broad (according to Hamet); inflorescence umbellate or paniculate, broad, 
with sparse, declinate, sparsely glandular -pubescent branches, leafy; 
flowers 9—18, small, with pedicels, ca. 1 mm long; sepals linear -lanceolate, 
acuminate, glabrous; petals pink, ovate -lanceolate, acuminate, entire, 
glabrous, often glandular -pubescent along the keel, 6—8 mm long, 3 times 
as long as calyx; stamens 10, the 5 opposite to petals half as long as and 
adnate to them nearly to the middle; pistils with short subulate -filiform 
style; hypogynous scales semiorbicular, entire or slightly emarginate, 
broader than long; follicles approximate, not inflated ventrally, erect, 
villous on the outside, 6— 7 mm long; seeds ovoid, less than 1 mm long. 

Caucasus: Recorded for S. Transc. — Armenia, Nakhichevan, Bezoglyban 
(Konig). This indication should be verified. Gen. distr.: Bal.-As.Min. 
Known from Asia Minor (Bithynia, Ankara, Karya). Described from 
Asia Minor. Type in Venice. 

Note. We have no specimen from the Caucasus. A species closely 
related to S. obt u s i fo li u m, distinguished by narrower leaves and by 
glandular -pubescent inflorescence branches (Grossheim). According to 
Hamet, the sterile shoots ofS.listoniae have semiterete instead of 
flat leaves and follicles are not divaricate; this species would thus be 
remote from S. obtusifoliu m, which is unlikely. Judging by the 
illustration ofS. cariense Jaub. et Sp., a synonym of S. 1 i s t o ni a e, 
this species is closely related to S.obtusifolium C. A. M. and to 
S. s t o lo ni f e r u m Gmel. Raymond-Ham et considers S. li st o ni a e Vis. 
as a synonym ofS. obtusifolium C.A. M., as well as of S. bornmulleri 
Hausskn. (S. r ho d a nt hum Bornm.), known from Mt. Deli -dag in Armenia. 



Subsection 2. CRASSIFOLIAE A. Bor.— Leaves alternate, terete or 
thickened, linear -subulate to oblong. 



Series 1. Albae Berger, 1. c, 452 (emend.).— Flowers white, pinkish, or 
^eddish. 

31. S.tenellum M. B., Fl. taur.-cauc. Ill (1819) 315; Ldb., Fl. Ross. II, 188 
Boiss., Fl. Or. U, 782; Hamet in Acta Horti Tiflis. VIII, 3 (1908) 19; 
Grossh., Fl. Cauc. 11,228; Berger in Engl. u. Pr., Nat. Pflzfm. 18a (1930) 
453.- Ic: Frod. in Acta Horti Gothoburg. VII (1932) pi. XVIH, textfig. 
212-220; Grossh. in Monit. Jard. Bot. Tiflis III, IV (1917) 135, f. 19. 

Perennial; roots fibrous; cespitose plants with flowering and sterile 
shoots, branching from base; stems numerous, glabrous; flowering 
shoots 4—10 cm long, erect or ascending, leafy, slender; sterile shoots 
1—3 cm high, with densely imbricated leaves; leaves alternate, oblong, 
terete, fleshy, free at base, with obtuse spur, sessile, glabrous, obtuse, 



63 



82 



entire, 3—5 mm long, ca. 1 — 1.5 cm broad, remote on fruiting stems, 
approximate on the sterile, green, sometimes reddish; inflorescence 
corymbiform or paniculate with few or up to 2 5 flowers, rather loose; 
pedicels as long or about as long as calyx; flowers small, 5-merous; 
corolla 1.5—2 times as long as calyx; calyx green, connate at base, 
with oblong-lanceolate obtuse lobes, glabrous; petals 5, glabrous, ovate - 
lanceolate or ovate, entire or slightly emarginate, apiculate, 3—4 mm long, 
red or whitish; stamens 10, 2 / 3 as long as petals, as long as or slightly 
shorter than pistils, with red filaments and dark reniform anthers; pistils 5, 
ovate or oblong-ovate, glabrous, with short curved style and capitate stigma; 
hypogynous scales large, about half as long as pistils, ca. 1 mm long, on long, 
erect pedicel, with an oblate, emarginate part ca. 1 mm broad, rounded or 
with a projecting at the top; follicle erect, connate below, convex in the 
upper part, on the inner side, near the beak; seeds oblong-ovoid, small, 
ca. 0.5 mm long, brown. June —July. 

Pebbly and stony soils in the high-mountain zone at 2000— 3000 m. — 
Caucasus: Dag., E., S., and W. Transc. Gen.distr.: Iran. (Elburz Mts.), 
Arm. -Kurd, (former Artvin District; Kars Region). Described from 
the Caucasus (Lars). Type in Leningrad. 

*32. S.lydiumBoiss., Diagn.pl. nov. ser. 1, 3 (1843) 17; Fl. Or. II, 782; 
Grossh., Fl. Cauc. II, 228; Berger in Engl. u. Pr., Nat. Pfzlfm. 18a (1930) 
453.- Ic: Praeger in Journ. Hort.Soc. (1921) f. 106; Frod. in Acta Horti 
Gothoburg. VII (1932) pi. XIX, textfig. 221-229. 

Perennial; roots fibrous, arising from a slender, creeping rhizome; 
cespitose plants with flowering and shorter sterile shoots, branching 
from base; stems ascending, (5)7— 10cm high, densely leafy, slender; 
leaves alternate, narrowly linear or lanceolate, terete, subobtuse, 4—6 (8) mm 
long, ca. 1 mm broad, glabrous, more approximate on sterile shoots, green 
or reddening; inflorescence dense, nearly capitate -corymbiform, ca. 1—2 cm 
broad, many -flowered, with short, densely flowered branches; pedicels 
shorter than or as long as calyx; flowers small, 5-merous; calyx nearly 
2 / 3 as long as corolla, with ovate, subobtuse sepals; petals pink, sometimes 
whitish, ovate -lanceolate, subobtuse, 3—5 mm long; stamens 10, scarcely 
shorter than petals, with white filaments and dark anthers; hypogynous 
scales yellow, elongate -cuneate-spatulate, emarginate; follicles erect 
white, later reddening, shorter than stamens, with slender beak ca. 1 mm 
long, minutely tuberculate; seeds glabrous, ovoid, subobtuse, very small, 
ca. 0.5 mm long. June. 

Moist soils in the subalpine zone.— Outside the USSR. Gen.distr.: 
Arm. -Kurd: former Artvin district; Bal. -As. Min: As. Min. Described 
from Asia Minor (Lydia, Karya). Type in Geneva. 

Note. Often in cultivation, used for carpet -like flower beds. 

33. S. album L., Sp.pl. (1753) 432; Ldb., Fl. Ross. II, 187; Boiss., Fl. 
Or. II, 781; Shmal'g., Fl. I, 362; Hamet in Acta Horti Tiflis. VIII, 3 (1908) 21; 
Grossh., Fl. Cauc. II, 228; Berger in Engl. u. Pr., Nat. Pflzfm. 18a (1930) 453.- 
Ic: Rchb., Ic. Fl. Germ. XXIII (1898-1899) tab. 55; Hegi, 111. Fl. IV, 2 (1925) 
tab. 140, f. 8; f. 916; Praeger in Journ. Hort. Soc. (1921 ) 182; Grossh. in 
Monit. Jard.Bot. Tiflis III-IV (1917) 135, f. 16; Frod. in Acta Horti Gothoburg. 
Vn (1932) po.XII-XIII, textfig. 151-160.- Exs. : Fl. Exs. Reipubl. Bohem. 
Sloven., No. 324; PI. Finl. exs., No. 250; HFR, No. 1717. 



64 



Perennial, with creeping rhizome and fibrous roots, cespitose in lower 
part, branching at base, with prostrate, sterile, densely leafy shoots, 2—3 cm 
long; stems fruiting, ascending; flowering shoots 8—20 (30) cm long, erect, 
slender, simple, glabrous; leaves alternate, terete or flattish, sessile, with 
a blunt short spur at base, remote from the stem, glabrous, oblong or 
oblong-ovate, obtuse, 7—10 mm long, 1—2 mm broad, remote on the 
flowering shoots and crowded on the sterile; inflorescence bifurcate - 
corymbiform or paniculate; pedicels as long or longer than calyx; flowers 
numerous, small, 5-merous; sepals glabrous, connate at base, broadly 
ovate, obtuse, green, persistent in fruit; petals white, ovate -oblong, 
entire, subobtuse, 3—5 mm long, 3—4 times as long as calyx, slightly connate 
at base; stamens 10, about as long as petals, with white filaments and red 
reniform anthers; stamens opposite to petals adnatetothemat base; hypogynous 
scales compact, small, broadly clavate, tapering to a short pedicel, convex 
at the apex or bluntly emarginate; follicles 5, erect, compressed, straight, 
pale green, not gibbous ventrally, oblong, with slender subulate beak ca. 1 mm 
long, villous inside; seeds small, oblong, ca. 1 mm long, brownish. June — 
July. 

Stony soils. European part : Crim. (?); Caucasus: mainly in E. Transc. 
and Cisc, less in S. and W. Transc. Gen. distr. : throughout Eur. from 
England and Scand. to Italy, Spain, Baltic States (Finland, Aland Islands, 
Hiiumaa, Saare), N. Afr., Bal. -As. Min., Arm. -Kurd. Described from 
Europe. Type in London. 

Note. Extremely polymorphic species needing further critical 
investigation. Dimensions of entire plant and of leaves, flowers, and 
inflorescence vary greatly. A whole series of species have been described, 
such as S. m ic r a nt hum Bast. (v. m i c r a nt hum DC.) with oblong leaves; 
S.turgidum Ram. (v. t u r gidum DC.) with ovate leaves; S.ath-oum DC, 
distinguished by acute petals (?); S.albellum Bess., S. b a It i cu m Hartm., 
and a number of other closely related species requiring thorough 
verification, and for the present recorded as synonyms of S. album. L. and Ldb. 
indicates S.transbaicalense Schlecht. pat. in Herb, reg. berol. as a 
synonym of S. album; this is undoubtedly an error, for S. album does not 
grow in Siberia. 

34. S.gracile C.A.M., Enum. plant. Cauc. (1831) 151; Ldb., Fl. Ross. II, 
186; Boiss., Fl. Or. II, 781; Hamet in Acta Horti Tiflis. -VIII, 3 (1908)25; 
Berger in Engl.u. Pr., Nat. Pflzfm. 18a (1930) 453.- S. albert i Praeger 
in Journ. Hort. Soc. (l92l) 191, non Rgl. — Ic: Frod. in Acta Horti Gothoburg. 
VII (1932) pi. XXX, textfig. 394-403; Praeger, 1. c, f. 107, 108; Grossh. in 
Monit. Jard. Bot. Tiflis. Ill and IV (1917) 135, f. 15. 

Perennial; roots fibrous, cespitose, branching plants with flowering 
shoots 4—6 cm long and with sterile shoots 1.5—3 cm long; flowering 
shoots curved or straight, ascending at base, slender, simple, glabrous, 
divaricate, densely leafy; sterile shoots densely imbricate -leaved; leaves 
alternate, 5—6 mm long, 1— 1.3 mm broad, terete, linear-subulate or linear - 
oblong, glabrous, sessile, subobtuse, with a remote blunt spur at the base; 
inflorescence corymbiform, broad, 1.5—2.5 cm long, many -flowered, leafy, 
with 2—3 elongated, declinate, secund, flexuous branches; flowers short - 
pediceled or sessile, small, numerous, 5-merous; bracts ovate, acute; 



65 



84 



sepals 5, connate a1 base, elliptic, broadly triangular, subobtuse, green, 
Y 3 — l /o as Long as corolla J petals 3— 4 mm long, lanceolate, acuminate, white, 
keeled; stamens 10, the 5 opposite to petals adnate to them to the middle; all 
stamens shorter than petals, as long as fruit, with white filaments and red 
anthers: hypogynous scales euneate, oblong, emarginate; follicles glabrous, 
ovoid-oblong with a beak 1—1.5 nun long, whitish or pale greenish, initially 
erect, later slightly divergent, convex on the inside; seeds small, brown, 
oblong-ovoid. June July. Stony soils in mountains in the subalpine and 
alpine .-.ones. — Caucasus: Cisc, Dag. (often); E. and W. Transc. (rarely); 
S. Transc. Gen. distr.: Arm. -Kurd., former Artvin District, 
Samachur Range. Described from the Caucasus, Gutgora. Type 
in Leningrad. 

35. S. lenkoranicum Grossh. in Monit. Jard. Bot. Tiflis. Ill— IV (1915) 
173; Grossh. Fl. Cauc. II, 229.- Ic: Grossh. in Monit. Jard. Bot. (1917) 
135, f. 19. 

Perennial, green, cespitose; stems ascending, (5) 10—15 (20)cm high, 
densely leafy, branching; sterile shoots densely covered with imbricate 
leaves: leaves alternate, terete, subulate, obtuse, 3— 6 mm long; inflorescence 
tew -flowered, subcorymbiform, 2— 3 -branched; flowers with 
bracts resembling cauline leaves, subsessile or with very short pedicels; 
sepals green, oblong-lanceolate, subobtuse, connate at base, nearly V 3 as 
long as petals; petals white or pinkish (especially when dry), lanceolate, 
acuminate, 6—7 mm long; stamens shorter than petals, slightly longer 
than pistils, with white filaments and dark anthers; hypogynous scales 
reduced, obtuse, emarginate; follicles lanceolate, convex, ca. 5 mm long, 
with filiform beak 1 mm long, not stellately spreading at maturity, erect; 
seeds light brown, small, ca. 0.5 mm long, elongate -ovoid. June — July. 
(Plate V, Figure 6a-b). 

Rocks in the high-mountain zone.— Caucasus: Tal. Gen. distr. : 
Arm. -Kurd, (former Kars Region), Iran. Described from mountainous 
Talysh. Type in Leningrad. 

Note. Species closely related to S. g r a c i 1 e C. A. M. but distinguished 
by its greater size, few -flowered inflorescence, larger flowers, and shape 
of hypogynous scales. In addition, the distribution area of this species 
does not coincide with that of S. gr a c il e, which is characteristic for the 
Greater and Lesser Caucasus. Meyer made a distinction between 
S. g r a c i 1 e with small and S. gracile with large flowers. 

36. S. subulatum (C. A. M.) Boiss., Fl. Or. II (1872) 783; Hamet in Acta 
Horti Tiflis. VIE, 3 (1908)21; Grossh., Fl. Cauc, II, 228; Frod. in Acta 
Horti Gothoburg. VII (1932) 23.- S.acutifolium Ldb., Fl. Ross. II 
(1844-1846) 187; Boiss., I.e.; Shmal'g., Fl. 1,362; Berger in Engl. u. Pr., 
Nat. Pflzfm. 18a (1930) 453.- S. calve rti Boiss., Diagn. Ser. 11,2 (1856) 
62.- Cotyledon subulata C. A. M., Enum. plant. Cauc. (1831 ) 150.- 
Umbilicus subulatus Ldb., 1. c, p. 173. — Ic: Grossh. in Monit. Jard. 
Bot. Tiflis III-IV (1917) 135, f . 1 . 



6o 



Perennial; roots thickened, numerous, cespitose. Plants with fruiting 
shoots and with densely leafy sterile shoots; stems little branching, 
suberect or ascending, (5)10—20 cm high, sometimes higher; leaves 
alternate, fleshy, subulate -linear, acuminate, spreading at base, spur red, 
erect, glabrous, glaucous -green, 9—11 mm long, ca. 1.5 mm broad, entire; 
inflorescence corymbiform, capitate, dense, with scorpioid branches; 
flowers with short pedicels as long as bracts, shorter than calyx, 5-merous, 
numerous; sepals glabrous, connate at base, ovate, acuminate, l / z — Y 2 
as long as corolla; petals oblong-ovate, erect, entire, obtuse or apiculate, 
white, connate at base, ca. 5 mm long; stamens 10, about as long as petals 
or slightly exserted, with white filaments and dark anthers; hypogynous 
scales very small, flabellate, broader than long, of dense tissue at center 
and of thin, transparent tissue on the margin, irregularly toothed; follicles 
glabrous, ovate -lanceolate, connate below, without gibbosity on the inside, 
with long subulate beak about as long as fruit without beak; seeds ovoid, 
brown, 1 mm long. June — July. (Plate V, Figure 7a). 

Stony soils, mountains, at 1 ,200— 2, 100 m. — European part : L.Don, 
(S. part of the Stalingrad [Volgograd] District near the village of Ivanovka — 
separated from the common distribution area) — Caucasus: Cisc. 
(Main Range, Voroshilovsk [Stavropol]), E. Transc, Dag., S. Transc, Tal. 
Gen. distr. : Arm. -Kurd. (Turkish Armenia, former Kars Region). Described 
from Talysh. Type in Leningrad. 

37. S. alberti Rgl., Descr.pl. nov. in A.H. P. VI, 2 (1879) 299. - S. a f f i n e 
Hamet in Candollea IV (1929-1931) 4, non Boreau (1866).- S. s c h r e n k i i 
Frod.in Acta Horti Gothoburg. VII (1932) 22. - Umb i 1 i c u s af finis 
Schrenk, Enum. pi. nov. I (1841 ) 72; Ldb., Fl. Ross. II, 173.- Cotyledon 
alfinis Maxim, in Mel. biol. XI (1883) tab. 23; Bull. Ac. Sc. Pet. XXIX 
(1884) 120. — Ps eudo s edu m affine Berger in Engl. u. Pr., Nat. 
Pflzfm. 18a (1930)465.- Ic: Frod., 1. c, pi. XI, textfig. 141-1 50; Gartenflora 
(1880) tab. 1019, f.2. 

Perennial; roots branching, numerous, cespitose, cordlike; stems 
numerous, ascending or erect, sterile, reduced, 1—5 (8) cm high, densely, 
almost imbricately leaved; fruiting stems 7—15 (25) cm high, mostly 
ascending at base, simple, the leaves arranged less compactly; leaves 
alternate, fleshy, subterete, rather flat in upper part, linear or linear - 
oblong, obtuse, surface covered with minute sharp tubercles, 2—7 (ll)mm 
long, 1 — 1 .5(2) mm broad at base with rounded or obtuse margin, ca. 0.5 mm 
broad, attenuate downward, and appressed to stem, inflorescence 
corymbiform, with scorpioid secund branches, 1.5— 2.5cm long, 2.5— 4 cm 
broad; flowers 5-merous, subsessile or with pedicels 0.5—1 mm long, shorter 
than calyx, sepals 1 .5—2 mm long, connate at base, ovate, tapering toward 
the apex, subobtuse, ca. 1 mm broad, l / 3 as long as corolla; petals white, 
ca. 5 mm long, 2 mm broad, connate only at base, oblong-ovate, apiculate, 
subobtuse; stamens 10, with dark violet anthers; the stamens opposite 
to sepals as long as or somewhat shorter than petals, those opposite to petals still 
shorter; hypogynous scales short and broad, crenulate-subtruncate, ca. 0.5 mm 
long; follicles many- seeded, oblong -ovoid, 3—3.5 mm long, 1.5 mm broad, with 
beak 2.5— 3 mm long, about as long as follicles, protruding from corolla; 
seeds small, oblong -ovoid, less than 1 mm long, brown, obtuse. Fl. May — 
July, fr. July — August. 



67 



Stony mountain slopes, rock crevices, pebbles, argillaceous bluffs, dry 
stony riverbeds, at ca. 1,100— 1,750 m. — W.Siberia: Irt. (Kokchetav, 
Pavlodar, and Karkaralinsk Mountains), Alt. (foothills of S. Altai); 
Centr.Asia: Ar. -Casp., (Ulu-Tau Mountains), T. Sh., Dzu. -Tarb. 
Gen. distr. : Dzu.-Kash. (Kuldja). Described from E. Turkestan. Type 
in Leningrad. 

Note. S.alberti Rgl. differs from U. aff i n i s Schrenk neither 
in its description nor according to the authentic specimen. The described 
species is closest to S. subulatum (C. A. M.) Boiss., from which it 
is distinguished by its linear -oblong leaves, subobtuse petals, subobtuse 
sepals, and completely different distribution area. 



Series 2. Rupestres Berger, I.e., 456.— Leaves linear -subulate, semi- 
terete, appendaged; flowers yellow. 

38. S. reflexum L., Sp. pi. ed.2 (1762)618; Shmal'g., Fl. I, 361 ; Grossh., 
Fl. Cauc. II, 227; Berger in Engl. u. Pr., Nat. Pflzfm. 18a (1930) 456.- 
Ic: Rchb., Ic. Fl. Germ. XXIII, tab. 60; Berger in Journ. Hort. Soc. (l92l) 
268. 

Perennial, with creeping rhizome; fruiting stems 15—25 (40)cm high, 
erect, ascending, creeping at base, not branching; sterile shoots creeping 
or ascending, branching; leaves greenish, glabrous, sessile, remote on 
fruiting shoots and crowded on the sterile, semiterete, linear-subulate, 
mucronate, with a small basal tubercle pointing downward; inflorescence 
corymbiform, initially dense, later rather loose, branching, the flowers 
short -pediceled, 5— 7-merous; sepals oblong -lanceolate, acute, green, 
fleshy, persistent in fruit, shortly connate at base; petals bright yellow, 
lanceolate or linear -lanceolate, acuminate, twice as long as calyx or 
longer, keeled on the back, divaricate; stamens 10— 14, yellow, shorter 
than petals; hypogynous scales yellow, quadrate; follicles yellowish, erect, 
as long as stamens, lanceolate, with long subulate beak; seeds small, 
numerous. July. 

Sandy and stony soils. — European part: M. Dnp. (Korystyshev — in 
gardens, probably introduced); Caucasus: reported for the upper mountain 
zone, on the slopes of the Terskei Ala-Tau (Grossheim, 1. c); there is a 
specimen from Kazbek in the herbarium of the Botanical Institute (collected 
by Lagovskii). Gen. distr. : Centr. Eur.: Poland, N., W., and Centr. Eur.; 
N.Am.— introduced. Described from Europe. Type in London. 

Note. The reports for the Caucasus are apparently erroneous. 
Ledebour's report concerning S ib i r i a uralensis (Lepekhin) and for 
the Inder Mountains (Pallas) should also be considered erroneous. 



Series 3. Mites Berger, 1. c, 455. — Leaves linear, terete, appendaged; 
flowers yellow. 

39. S. sexangulare L., Sp. pi. (1753) 432; Praeger in Journ. Hort. Soc. 
(1921) 263; Frod. in Acta Horti Gothoburg. VII (l93l) 60; Pachoskii, Fl. 
Poles'ya in Tr. Khar'k. O-va Estestv.,XXX (1900) p. 81; Huber in Fedde, 



68 



Repertorium sp. nov. XL, 20-25 (1936) 364.- S. mite Gilib., Fl. Lithuan. V 
(1781)192; Hegi, 111. Fl. Mittel-Europa, IV, 2 (1925)539; Berger in Engl, 
u. Pr., Nat. Pflzfm. 18a (1930)455.— S.boloniense LoisL in Desv., Journ. 
Bot.,2 (1809) 327; Shmal'g., Fl. I, 361; Pachoskii, I.e., XXVII, 208.- Ic: 
Rchb., Ic. Fl. Germ. XXIII, tab. 57; Praeger in Journ. Hort. Soc. (1921 ) 
f. 155; Frod., in 1. c., pi. XXXV, textfig. 478— 496; Hegi, I.e., tab. 140, f. 5, 
f. 920. 

Perennial; roots slender, cordlike, arising from creeping, branching 
rhizome; stems 8—18 cm high, creeping, branching, ascending, densely 
leafy, fruiting shoots overtopping the sterile; leaves linear, terete, with 
an obtuse basal appendage, subobtuse, ca. 0.5 cm long, ca. 1 mm broad, 
more crowded on sterile shoots; inflorescence a loose, corymbiform 
cyme, usually with 3 (4)branches; bracts ovate, acute; flowers sessile, 
5-merous; sepals green, linear or linear -lanceolate, connate at base, 
ca. 2 mm long, obtuse; corolla twice as long as calyx, pale yellow, with 
linear-lanceolate, acute petals, ca. 4 mm long, ca. 1 mm broad; stamens 10, 
shorter than petals, yellow; hypogynous scales yellow, small, subquadrate; 
follicles as long as stamens, erect, with long filiform beak; seeds small, 
numerous, rounded, obtuse. June— July. 

Dry sandy and stony soils.— European part : S. sexangula re may 
occur in the USSR, but only in the Far West; reported for M. Dnp. (Podolia, 
near the Zbruch, Dniester, and Bug rivers, near Goloskov) (Shmalhausen). 
Gen. distr. : Poland and farther west, almost all of Europe; occurs in the 
Aland Islands; naturalized in some sites in England. Described from 
N. Europe. Type in London. 

Note. Many authors have considered S.sexangulare as one of the 
forms of S. ac r e (Hegi, Berger), or as its synonym; often the same 
illustrations were labeled both as S. a c r e and as S. sexangulare (for 
example in Maevskii in old editions, in Syreishchikov, etc.). Judging 
by Linnaeus 's short description, S.sexangulare can be considered as 
one of the ecological forms of S. a c r e. However, basing our decision 
on the work of Huber (l. c), who has seen authentic specimens from the 
Linnaean Herbarium, we follow the example of Praeger (l. c.) and 
Froderstrom (l. c.) in considering S. sexangulare L. the same as 
S. mite Gilib., but S. bo 1 o ni e n s e Lois, an independent species. 



Series 4. Acres Berger., 1. c, 454. — Leaves triangular -ovate, broadening 
toward base, unappendaged. Flowers yellow. 

40. S.acre L., Sp.pl. (1753) 432; Ldb., Fl. Ross. II, 187; Boiss., Fl. Or. II, 
783; Shmal'g., Fl. I, 361; Kamch., Fl. Zap. Sib. VI, 141 3; Berger in Engl, 
u. Pr., Nat. Pflzfm. 18a (1930)454.— S. sexangulare auct., non L. — 
Ic: Rchb., Ic. Fl. Germ. XXIII, tab. 51; Borisova in Sbornye rast. SSSR 
III (1934) 115; Frod. in Acta Horti Gothoburg. VII (1932) pi. XXXV, textfig. 
516-526.- Exs.: PL Finl. exs. No. 701 ; Fl. polonica exs. No. 634; HFR 
No. 462, 

Perennial, glabrous, with slender cordlike, creeping and branching 
rhizome; stems numerous, ascending or decumbent, fleshy; flowering shoots 
4—15 cm high, ascending; sterile shoots 1—3 cm high, often decumbent; 



69 



leaves fleshy, ovate, obtuse, convex on the back, thickened at base, with 
rounded gibbosity, 2—5 mm long, 2 — 3.5 mm broad, imbricatc-'y arranged 
in 5 or 6 rows on sterile shoots, more remote on flowering shoot" - 
inflorescence a cyme of 3—5 spiciform branches, spreading or compressed, 
the flowers subsessile, 5-merous; sepals 2— 3 mm long, oblong-ovate, 
glaucous -green or yellowish, free to the base; corolla 2—3 times as long 
as calyx, with golden-yellow, lanceolate or linear -lanceolate, acute petals 
4.5 mm long, 1.5—2 mm broad; stamens 10, slightly shorter than petals; 
hypogynous scales broader than long or subquadrate, ca. 0.5 mm long and 
broad; follicles pale green or whitish, 3.5—4.5 mm long, lanceolate, stellately 
spreading, with short straight beak; seeds small, numerous, 0.5 — 1 mm long, 
ovoid, light brown. May— July. 

Dry elevations, mostly on sandy soil, shore sands, sometimes stony 
and limestone soils; sometimes as a weed, in crop fields, mainly at the 
edges. Widespread almost throughout the RSFSR* from the Kola Peninsula 
and the Solovetskie Islands to the Crimea and the Caucasus; in W.Siberia 
from 60.5° N. (rare). — European part : Kar. -Lap., Lad. -Ilm., Dv. -Pech. — 
common in W. Perm Region, rare in E. part (Cherdyn, Verkhotur'e), 
V.-Don, BL, L.Don, Transv., L. V., Crim.; W.Siberia: reported for Ob, 
near the village of Zyranka on the Pyshma River and near Lake 
Alexandrovskoe (Schmalhausen); Caucasus: Cisc, W. and E. Transc. 
Gen. distr. : Bal.-As. Min., Med., nearly all of W.Eur., with the exception 
of the southernmost part of islands in southern Italy and most of the 
Balkan Peninsula, As. Min., N. Afr., N. Am. — introduced. Described from 
Europe. Type in London. 

Economic importance. The sap of the green parts of S. ac r e is 
poisonous; used locally to provoke blushing (the "water of life" of folk 
tales); freshly collected plants applied to the skin cause inflammation and 
blisters, hence the vernacular name "pryshchinets" (pimple); it has the 
same effects as Ranunculus flammula. 



Series 5. Japonicae Maxim, in Bull. Acad. Pet. XXIX (1884) 148.— 
Leaves linear, without basal appendage; flowers yellow. 

*41. S. polytrichoides Hensl. ex Forbes et Hemsl. in Journ. Linn. Soc. 
Lond. XXIII (1886-1888) 186; Kom., Fl. Manch. II, 398. - S. c o r e e n s e Nak. 
in Fedde, Repert. XIJI (1913-1915) 272.- Ic: Forbes et Hemsl., 1. c, 
tab. VII B, f . 4 ; Frod. in Acta Horti Gothoburg. VI (l 930 ) pi. LV, textf ig. 
694-702. 

Perennial, cespitose, with slender creeping rhizome, covered by dark 
brown dry leaf remnants; stems (3)5— 10cm long, numerous, branching 
from base, some sterile, slender, curved, ascending, densely leafy; leaves 
alternate, narrow, lanceolate -linear or linear, without basal appendage, 
more remote on flowering shoots, 0.5—1 .5 mm long, 1—2 mm broad; 
inflorescence corymbiform, 2— 3 -branched; flowers 3— 15, short -pediceled, 
stellately open, 5 -merous; sepals short, %— l / 3 as-long as corolla, ovate, 
1.5—2 mm long, acuminate; petals nearly free, narrowly lanceolate, golden- 
yellow, acuminate, 5—6 mm long; stamens shorter than flowers, with yellow 






* [The Russian Soviet Federated Socialist Republic] 



70 



filaments and rounded anthers; hypogynous scales small, suborbicular, 
slightly narrowing toward base; follicles substellately spreading, with 
long, erect beak, ca. 1.5 mm long, connate at base, 4.5—5 mm long, ovoid- 
oblong; seeds oblong, small, less than 1 mm long, brown, obtuse. Fl. 
July— August, fr. August— September. 

Forests, moist rocks covered by moss carpets, shady or somewhat 
shady sites; growing in large groups. At USSR borders. Gen. distr. : 
Jap.-Ch. (Kirin Province, Manchuria). Described from Chekiang, Ningpo. 
Type in London. 



Section 5. EPETEIUM Boiss, Fl. Or. II (1872) 776.- Flowers 5-merous, 
less often 4— 9-merous; inflorescence corymbiform or umbellate, with 
2 to few branches, or spiciform. Mainly annuals, less often biennials, with 
slender roots; flowering specimens without sterile shoots; leaves 
alternate, semiterete or terete, less often flattish. Species distributed 
mainly in the Mediterranean area. 



Series 1. Hispanicae A.Bor.— Stamens in two series, twice as many as 
petals. Calyx shorter than petals. 

42. S. hispanicum L., Sp. pi. (l 753) 61 8, non S. h i s p a n i c u m Raymond 
Hamet; Ldb., Fl. Ross. II, 185; Shmal'g., Fl. I, 362, Berger in Engl. u. Pr., 
Nat. Pflzfm. 18a (l 930 ) 46 1 ; Frod. in Acta Horti Gothoburg. VII (l 932 ) 79. - 
S. glaucum Waldst. et Kit., Descr. et icon, plant, rare. Hungar. II (1805) 
198; Boiss., Fl.Or. II, 789.- S. sexf idum M. B., Fl. taur.-cauc. I (1808) 
354.— S.orientale Boiss., Diagn. Ser. I, 10 (1849) 17. — S. a rme num 
Boiss., Diagn. Ser. II, 2 (l 856) 61.— S.heptapetalum Fisch. in Ldb., 1. c, 
186.- Ic: Rchb., Ic. Fl. Germ. XXIII, tab. 51 ; Waldst. et Kit., 1. c, tab. 181; 
Sibthorp, Fl. Graeca 5, tab. 449; Jacquin, Fl. Austriac. 5, tab. 47; Hallier, 
Fl. Deutschland 26, p. 2643; Praeger in Journ. Hort. Soc. (1921 ) f. 178; Hegi, 
111. Fl. f. 912; Grossh. in Monit. Jard. Bot. Tiflis. Ill- IV (1917) 135, f. 20.- 
Exs. : Fl. exs. Austr. -Hung., No. 3680. 

Annual or biennial; roots short, little developed; stems usually branching 
from base, 5 — 10 (20)cm long, more or less glandular -pubescent, with 
ascending, simple branches; leaves sessile, alternate, linear, or oblong- 
lanceolate, flat or semiterete, mostly subacute, slightly tapering toward the 
apex, glabrous, pale green, often reddening, 0.5—0.8 cm long, diminishing 
toward the inflorescence; inflorescence of 3—4, often secund branches, 
leafy; flowers short -pediceled, or subsessile, 6 -, sometimes 7— 9-merous; 
sepals pubescent, ovate, acute, Y 3 as long as corolla, green; petals lanceolate, 
acuminate, glabrous, white with a reddish median line, subcarinate, 4—5 mm 
long; stamens 12 (14—18), shorter than petals, longer than pistils, with 
white filaments and subrounded dark purple anthers; pistils compressed - 
conical, initially white, later reddening, pubescent, with glabrous subulate 
style; hypogynous scales yellow, subternate, narrower at the base; follicles 
6 (7—9), stellately spreading, rounded -ovoid or ovoid, ca. 2 — 3 mm long, 
with a ventral protuberance arising at very tip of fruit from subulate, 
somewhat curved beak; seeds numerous, very small, oblong-ovoid, 



71 



91 



compressed, subobtuse, brown. June — July. (Plate V, Figure 2). 

Dry stony slopes, mainly in the middle zone of mountains.— European 
part: Crim.; Caucasus: mainly E. Transc. (Main Range), Cisc, Dag., less 
in W. and S. Transc. and Tal. Gen. distr. : Arm. -Kurd, (former Artvin 
District), Med., Bal. -As. Min. Described from Spain (?). Type in London. 

Note. Linnaeus 's report on the occurrence ofS.hispanicum in 
Spain is doubtful; however, the name he gave has priority and must be 
retained. It is most probable that this species has been described from 
specimens grown in botanical gardens. 

43. S. pentapetalum A. Bor. sp. nova in Addenda VIII, p. 367. — 
S. h i s p a n i c u m auct. — S. g 1 a u c u m auct. 

Annual, not branching, with erect stem, with few erect branches only 
in the inflorescence; stems solitary, leafy, more or less glandular- 
pubescent, erect, 5—10 (l5)cm high; leaves sessile, oblong -lanceolate, 
fleshy, subobtuse, sparsely glandular -pubescent, 1—2 cm long, 1— 3 mm 
broad; inflorescence corymbiform, leafy, of (l ) 2 — 3 branches arranged 
in upper third of the plant; inflorescence branches usually secund, many- 
flowered, sometimes 1—2 flowered; flowers 5-merous, subsessile or short - 
pediceled, with bracts resembling cauline leaves: sepals green, sparsely 
glandular -pubescent, oblong-triangular, acute, x / 4 — Y 3 as long as corolla; 
petls oblong or linear -lanceolate, acuminate, glabrous, white or pinkish, 
with green median line, drying pale yellow, 5—6 (7) mm long; stamens 10, 
shorter than petals, with white filaments and dark, rounded anthers; 
pistils lanceolate, shorter than petals, with erect, subulate style; hypogynous 
scales small, tapering toward base, broadening at the summit, 2— 3-lobed; 
follicles stellately spreading, oblong -lanceolate, flat, with inconspicuous 
ventral protuberance, with erect subulate beak at base of fruit, minutely 
tuberculate on the surface, sparsely glandular -pubescent ventrally, less 
so dorsally, 5— 6 mm long; seeds ovoid, very small, obtuse. April — May. 
(Plate V, Figure la). 

Stony and pebbly mountain slopes, at 1,000— 2,000 m (species associated 
with mountain -xerophytic zone).— Caucasus: mainly E. Transc. (Kirovabad, 
Agdam Nukha, Kuba, Shusha, Geokchai), Dag. (Derbent), S. Transc. (Erevan), 
(solitary near Borzhomi), Tal. (between Lerik and Buzachar); Centr. Asia: 
Mtn. Turkm. (Bol'shoi Balkhan and Kopet Dagh ranges). Gen. distr. : 
Arm. -Kurd. (Kars Region), Iran, (there are specimens from the Elburz). 
Undoubtedly occurring in parts of Iran bordering on Kopet Dagh and the 
Caucasus. Described from Turkmenistan, Kopet Dagh Range. Plant 
of the Temperate Zone. Type in Leningrad. 

Note. This species is clearly differentiated from S.hispanicum L. 
by its simple, erect stem, not branching from the base, 5-merous flowers, 
shape of fruit, larger flowers and fruit. In addition, in S. pentapetalum 
anthesis occurs much earlier (April — May) than in S.hispanicum 
(June — July). The distribution areas of these 2 species also differ: 
S.hispanicum L. is an E. Mediterranean and European element, 
S. p e nt a p e t a lu m A. Bor.,. a typical Asiatic xerophyte associated with 
the mountain-xerophyte zone of the E. and SE Caucasus, Iran, and Mtn. Turkm. 
Frod. (in Act. Hort. Gothoburg. VII, 3 (1932) 82) quotes S.hispanicum var. 
semiglabrum (S. semiglabrum Boiss. et Huet in herb. ) from 



72 



Asia Minor, also with 5-merous flowers, but differentiated from our species 
by a series of characters: glabrous leaves, size of leaves, shape of sepals 
and hypogynous scales, the ovate petals pubescent along the midnerve. 

44. S. bucharicum A. Bor. sp. nova in Addenda VIII, p. 367. 

Annual, 4—5 cm high, branching, with numerous slightly curved, leafy 
branches forming a corymbiform inflorescence in the upper part, or else 
not branching, the inflorescence 2 -branched; stems solitary, leafy, 
sparsely glandular-pubescent, mainly in upper part; leaves sessile, 
alternate, linear, ca. 0.8 cm long, ca. 1 mm broad, subobtuse, with sparse 
glandular pubescence confined to lower part of leaves; inflorescence 
corymbiform, consisting of a series of branches usually bifurcate in 
upper part, with unilateral flowers; flowers 6-merous, short-pediceled 
or subsessile, with bracts resembling cauline leaves; calyx connate at 
base, consisting of 6 oblong, triangular, acute sepals with sparse hairs, 
2 / 5 as long as corolla, green; petals glabrous, lanceolate, acuminate, white 
with inconspicuous midnerve, 4 mm long, ca. 1 mm broad; stamens 12, shorter 
than petals, with dark anthers and white filaments; pistils with erect, 
filiform style 1 mm long, lanceolate, shorter than petals; follicles 6, 
stellately spreading, oblong -triangular, gradually broadening toward base, 
flatfish, 4 mm long, minutely tuberculate, sparsely short glandular-pubescent; 
seeds small, ca. 0.5 mm long, ovoid, acute on one side, obtuse on the other, 
brown. Fr. June. 

Mountain limestones.— Centr. Asia: Pam. -Al. (Sarsaryak Range, near 
the village of Tashanur; Vakhsh River, Sangtuda village). Endemic. 
Described from the Sarsaryak Range in E. Tadzhikistan. Type in 
Leningrad. 

Note. S. bucharicum A. Bor. is the eastern species, replacing 
other species of the series, Hispanicae A. Bor. It is differentiated 
from S.hispanicum by the shape of its growth, by larger — 4 mm long — 
follicles oblong-triangular and gradually broadening toward base, by a 
series of other morphological characters, and by its distribution area. 
The other closely related species, S. pentapetalum A. Bor., differs 
from S. bucharicum A. Bor. in the following characters : flowers 
5-merous; petals 5—6 (7) mm long; follicles 5— 6 mm long, lanceolate, 
with basal protuberance ventrally; stems erect, not branching; distribution 
area: Caucasus and Mtn. Turkm. 

45. S. pallidum M. B., Fl.taur.-cauc. I (1808) 353; Ldb., FL Ross. II, 
185; Boiss., FL Or. II, 790; Shmal'g., FL I, 362 ; Berger in Engl. u. Pr., Nat. 
Pflzfm. 18a (1930) 461. - S.hispanicum Hamet in Acta Horti Tiflis 
VIII, 3 (1908) 30, p.p., non L.; Frod. in Acta Horti Gothoburg. VII (1930) 

79p. p. — C r a s sula rubens var.decandra DC, Prodr. Ill (1828) 405. — 
Ic: Grossh. in Monit. Jard. Bot. Tiflis. III-IV (1917) 135, f. 11.- Exs.: Herb. 
Fl. Cauc, No. 322. 

Annual or biennial; stems 5—20 (30)cm high, usually branching from base, 
with weak, ascending, arched -upcurved branches, with glandular pubescence 
mainly in upper part on inflorescence branches; leaves alternate, oblong- 
lanceolate, semiterete, obtuse, sessile, declinate, 0.5 — 1 cm long; inflorescence 
branches 2—4, arched -upcurved, glandular -pubescent, many -flowered, 
93 secund; flowers 5-merous, subsessile, 3 mm long; calyx 2 / 3 as long as 



73 



94 



corolla, connate at base; petals pink, with dark midnerve, lanceolate, 
aristate -acuminate, erect; stamens 10, shorter than petals, with white 
filaments and dark anthers; hypogynous scales oblong, longer than broad, 
uneven at the summit; follicles obliquely ascending, not stellately 
spreading, with small ventral protuberance, usually glandular -pubescent, 
ovoid, with long straight beak; seeds very small, ca. 0.5 mm long, ovoid, 
brown. May — July. (Plate V, Figure 3a). 

Mainly in lower part of mountains, rising to 1,800 m; adapted to moist 
sandy and stony soils.— European part: Crim.; Caucasus: mainly W., E., 
and S. Transc, Tal., less in Cisc, Dag. Gen.distr.: Bal. -As. Min. ; 
Arm. -Kurd., Iran. Described from the Crimea. Type in Leningrad. 

Note. S. pallidum M. B. has often been confused with 
S.hispanicum L. Hamet and Froderstrom unite these species, although 
they are very well differentiated. Both species are widely distributed 
through the Caucasus, but S. pallidum occurs in moister and lower 
sites than S. hispanicu m. 

46. S. corymbosum Grossh. in Monit. Jard. Bot. Tiflis. III-IV (1915) 171; 
Grossh., Fl. Cauc. II, 230.- Ic: Grossh., I.e., III-IV (1917) 135, f. 8. 

Annual, light green; stems 5 — 15 cm high, ascending, glandular mainly 
on inflorescence branches, branching from middle of plant or higher, or 
else branching from base, the branches ascending at an acute angle; 
leaves terete, thickish, obtuse, glabrous or sparsely glandular, to 15 mm 
long, diminishing toward inflorescence; inflorescence a regular corymb; 
pedicels glandular, erect, especially in fruit, 3—4 mm long, slightly shorter 
than or as long as fruit; flowers usually 5-, less often 3— 6-merous, 
3—4—4.5 mm long; calyx 1 / 4 — l / 3 as long as corolla with triangular, acute, 
glandular sepals; petals whitish-greenish, acute; stamens 10 (6 — 12); 
hypogynous scales cuneate, furcate, with one or two acute teeth often as 
long as scales at the summit and several smaller teeth; follicles connate 
to Y 3 — Y 2 , erect, narrowly lanceolate, glaucous, often reddish -glaucous, 
4.5—5 mm long, glandular on the inner margin; seeds small, oblong, light 
brown. May— June. 

Adapted to dry stony slopes.— Caucasus: E. Transc, Dag., S. Transc. 
(Arak River valley between Negram and Daroshan), Tal. (Zuvant). 
Gen.distr.: possibly in N.Iran. Described from the Caucasus. Type in 
Leningrad. 

47. S.atratum L., Sp. pi. ed.2 (1763)1674; Boiss., Fl. Or. II, 792; Grossh., 
Fl. Cauc. II, 230; Berger in Engl. u. Pr., Nat. Pflzfm. 18a (1920) 460.-. 

Ic: Coste, Fl. Fr. f. 1363; Hegi, 111. Fl. IV, 2, tab. 141, f. 4, f. 913; Rchb., Ic. 
Fl. Germ. XXIII, tab. 153; Frod. in Acta Horti Gothoburg. VII (1932) pi. XLIX, 
textfig. 694-703.- Exs.: F. Schulz, herbarium normale, No. 662 et 662 bis. 

Annual; stems simple or branching from base, erect or ascending, 
with simple branches, 3—8 cm high, glabrous; entire plant usually dark or 
brownish-purple, densely leafy; leaves alternate, obtuse, ovate or oblong, 
clavate -terete, fleshy, sessile, approximate; inflorescence terminal, 
few-flowered, compact, corymbiform, leafy; flowers about as long as 
pedicels, 5-merous; sepals triangular -ovate, ca. 2 mm long, acute, connate 
at base, reddish; petals elongate -ovate, acuminate, 3—4 mm long, twice as 



74 






long as calyx, whitish-greenish or greenish pink; stamens 10, shorter than 
petals, ca. 2.5 — 3 mm long, with yellow reniform anthers and white filaments; 
hypogynous scales small, subquadrate; follicles five, 3— 4 mm long, ovoid, 
obtuse, with short filiform style, ca.0.5 mm long; seeds elongate -ovoid, 
very light brown, 0.5— 0.8 mm long. June — August. 

Stony soils, at 650—2,600 m. — Caucasus : E. Transc. (in upper part of 
Chengil-Chai — Lagovskii). Very doubtful indication — confusion of labels 
is possible. Gen. distr. : European mountains: Pyrenees, Alps, S. Jura, 
Carpathians, W. Tatra Mountains, Apennines; Bal. -As. Min. : mountains of 
the Balkan Peninsula. Described from Italy. Type in London. 

48. S.villosum L., Sp.pl. (1753) 432; Shmal'g., Fl. I, 363; Grossh., Fl. 
Cauc. II, 230; Berger in Engl. u. Pr., Nat. Pflzfm. 18a (1930) 461.- Ic. : 
Praeger in Journ. Hort. Soc. (1921 ) f. 179; Rchb., Ic. Fl. Germ. XXIII, tab. 52; 
Frod. in Acta Horti Gothoburg. VII (l 932) pi. XX, XXI, textfig. 264-269. 

Biennial; stems 4 — 15 cm high, erect or ascending, slightly branching 
or simple; entire plant glandular -pubescent; leaves linear -oblong, 3 — 5 mm 
long, obtuse, semiterete, flab above, inflorescence a simple or branching 
raceme, few-flowered; flowers 5-merous; pedicels longer than flowers; 
sepals obtuse, oval, glandular-pubescent; petals ovate, acute, 3—5 mm long, 
pink, darker below along keel, 2— 3 times as long as calyx; stamens 10, 
shorter than petals, with purple anthers; hypogynous scales spatulate- 
quadrate; follicles ovoid, with filiform erect beak; seeds numerous, ovoid, 
obtuse, small, ca. 1 mm long. June — July. 

Peaty meadows and marshes in the alpine zone. — Reported for the 
Caucasus — former Terskei Region (Grossheim). The herbarium of the 
Botanical Institute has a specimen from the Kazbek (collection of Lagovskii! 
Apparently, the report for the Caucasus is erroneous. Gen. distr. : N. and 
Centr. Eur., W.Greenland. Cited for Lithuania, Poland, and the Baltic 
States (near Riga — Shmalhausen). Described from Europe. Type in 
London. 

49. S.annuum L., Sp.pl. (1753) 432; Ldb., Fl. Ross. II, 186; Boiss., 
Fl. Or. II, 792; Hamet in Acta Horti Tiflis. VIII, 3 (l 908) 33; Grossh., Fl. 
Cauc. II, 229; Berger in Engl. u. Pr., Nat. Pflzfm. 1 8a (1930) 462.- 

S. saxatile DC, Fl. Fr. IV (1805) 394; DC, Prodr. Ill (1828) 409.- 
Ic: Rchb., Ic. Fl. Germ. XXIII (l 898-1 899) tab. 54; Hegi, III, Fl. IV, 
2 tab. 139, f. 8; Grossh. in Monit. Jard. Bot. Tiflis III-1V (1917) 135, 
f.21; Praeger in Journ. Horti. Soc. (l 921 ) f.182; Frod. in Acta Horti 
Gothoburg. VII (1932) pi. XXXII-XXXIV, textfig. 429-456. - Exs.: 
Fl. Exs. austro-hung., No. 2567; Fl. stiriaca exs. A. v. Hayek, No. 122 ; 
F. Schultz, herb, normale cent. 7, No. 663; PL Finl. exs., No. 251 et 
No. 702; Herb. Fl. Cauc, No. 320. 

Annual; roots fibrous. Plant branching from base or simple, 
glabrous, 4—1 5 cm high; sterile shoots absent; leaves alternate, 
terete or flatfish, convex on both surfaces, fleshy, sessile, glabrous at 
base, oblong-linear or ovate -oblong, 3— 6 mm long, 1—2 mm broad, 
obtuse, entire, pale green; inflorescence broad, leafy, consisting of 



75 



ramified branches covered with flowers and with flowers in 
bifurcations; flowers numerous, small, with pedicels as long as calyx 
or the flowers subsessile; sepals glabrous, connate at base, oblong- 
ovate, resembling the leaves, fleshy, obtuse, green; petals yellow, 
glabrous, twice or nearly twice as long as calyx, entire, apiculate, 
ovate -lanceolate, 3—4 mm long; stamen 10, slightly shorter than 
petals, with yellow anthers; the 5 stamens opposite to petals, connate 
to them nearly to the middle; hypogynous scales oblong, tapering 
toward base, bilobate at the summit; follicles 5, glabrous, ovoid - 
oblong, as long as stamens, greenish yellow, with filiform, erect 
style, initially erect, later declinate; seeds oblong, small, brown. 
May — June. 

Rocky sites in high-mountain regions (Arctic -alpine species).— 
Caucasus: Mainly E. Transc, Dag., less often S. Transc, Tal., 
W. Transc, Cisc. Gen. distr. : Scand., Centr. Eur., Med., Greenland, 
Bal. -As. Min., Arm. -Kurd., N. Iran. Described from N.Eur. Type 
in London. 

*50. S.nanum Boiss., Diagn.pl. nov.Ser. 1, 6 (1845) 57; Fl. Or. II, 
794; Grossh., Fl. Cauc. II, 237; Berger in Engl. u. Pr., Nat. Pflzvm. 
18a (1930) 461.- Ic: Frod. in Acta Horti Gothoburg. VII (1932) 
textfig. 254-263. 

Annual; roots fibrous, few; stems glabrous, 2—5 (10) cm high, 
erect, simple, weak; leaves lanceolate to ovate, semiterete, subacute, 
g g 3— 6 (8) cm long; inflorescence (l— 2) 5— 15 -flowered, paniculate, loose, 
flowers small, 5 -merous, long -pediceled, exceeding perianth; sepals 
ovate -triangular, acute, ca. 1 mm long; petals oblong, acute, 2.5— 3 mm 
long, 3—4 times as long as calyx, yellow, sometimes reddening along 
the keel; stamens 10, of which 5 about as long as petals, the other 
5 shorter; hypogynous scales s pat ulate -quadrate, emarginate; 
follicles ovoid, 2— 2.5 mm long, with short beak; seeds oblong, small, 
glabrous. June — July. 

Alpine zone, in the border zone beyond the USSR: Iran, (found 
not far from the Transcaucasian border and in S.Iran); Arm. -Kurd. - 
near Erzerum at 2,700 m. Described from S. Iran. Type in 
Geneva. 

Note. May occur in the Transcaucasian mountain ranges; 
being a very small plant, it may easily have been overlooked. 



Series 2. Rubrae A.Bor.— Stamens uniseriate, as many as petals; 
sepals shorter than petals. 

51. S.rubens L., Sp.pl. (1753) 432; DC, Prodr. Ill, 405; Berger 
in Engl. u. Pr., Nat. Pflzfm. 18a (1930) 461.- Crassula rubens L., 
Syst. Nat. ed.X (1759) 969.- Aithales rubens Webb, et Berth. 
in Phyt.Canar.I (1836) 179.- Ic: Rchb., Ic. Fl. Germ. XXIII (1898-1899) 



5773 76 



tab. 142; Hegi, 111. Fl. Mit. Eur. tab. 139, f. 9, f. 902 ; Praeger in Journ. 
Hort. Soc. (1921) f. 181 (except the flower analysis, with incorrect 
number of stamens); Frod. in Acta Horti Gothoburg. VII (1932) 
pi. XLV, XLVI, textfig. 600-614. 

Annual, rarely biennial; stems 5 — 15 cm high, erect, usually branching, 
less often simple, glandular -pubescent, viscous; leaves fleshy, semiterete, 
oblong -linear, obtuse, sessile, spreading at base, flat above, rounded below, 
reddish, remote; inflorescence consisting of simple, leafy, secund branches 
(spikes) with sessile, solitary flowers; bracts ovate -lanceolate; flower 
buds ribbed, hairy, red along the ribs; flowers 5-merous; calyx glandular - 
pubescent, connate at base, with broad, subovate, acute sepals, green or 
reddish, fleshy; petals broad -lanceolate to ovate, acuminate, white or 
reddish, with purple keel, 4—6 mm long, 3—4 times as long as sepal, hairy 
on outer side; stems 5, slightly shorter than petals, with white filaments and 
red anthers; hypogynous scales spatulate -quadrate, white, small; follicles 
initially erect, later divergent, lanceolate, acuminate, 4—5 mm long, minutely 
tuberculate and glandular -pubescent or glabrous, white, or reddish, 
compressed, as long as stamens. May — July. 

Fallows, plowland, roadsides. — European part: Crim., reported by 
Shmalhausen for southern coast near Nikita and Magarach; this species 
may have been confused with S. pa Hi d um. In Prodr. Ill, 405, DC. cites 
for the Crimea var. decandrum, with 10 stamens, related to 
S. pallidum M. B. Gen.distr.: Eur. and Med., S. and Centr. Eur., N. Afr., 
Canary Island; Iran.: N.Iran; Bal. -As. Min. : As. Min. Described from 
Europe. Type in London. 

Economic importance. Ornamental plants, suitable for carpetlike flower 
beds. 

52. S.rubrum (L.) Thell. in Fedde, Rep. spec. nov. X (1912) 290; Berger 
in Engl. u. Pr., Nat. Pflzfm. 18a (1930) 461; Frod. in Acta Horti Gothoburg. 
VII (1932) 89.- S.caespitosum DC, Prodr. Ill (1828) 405.- S. desert i - 
hungarici Simonkai in Oesterr. bot. Zeitschr. XL (1890) 333. — 
S. melanoleucum Schlecht. pat. in herb. reg. berol. ex Ldb., Fl. Ross. II 
(1844— 1846) 172. — C r a s s ula caespitosa Cav., Ic. et descr. pi. Hisp. I 
(1791)50; Ldb., I.e.; Shmal'g., I, 360. - C. m a g no 1 i i DC, Mem. Soc. 
Agric. Paris (1808) 11; Ej., Fl. Franc. VI (1815) 522. - T i 1 1 a e a rubra L., 
Sp. pi. ed. I (1753) 129. — T. caespitosa Four, in Ann. Soc. Linn. London 
N.S.XVI (1868) 384.- Procrassula magnolii Griseb., Spicil. Fl. 
Rumel.I (1843) 323.- Aithales caespitosa Webb, in Webb, et Berth., 
Phyt.Canar. I (1836) 179.- Ic. : Cav., I.e., tab. 69, f. 2; Rchb., Ic. Fl. Germ. 
XXIII, tab. 43; Frod., 1. c, pi. L., 2-3, textfig. 712-725. - Exs. : Herb, normale 
a T. Dorfler editum. No. 5265 (sub C r a s s u la caespitosa Cav.). 

Annual, subglabrous, 2—6 cm high; stem simple or little branching from 
base; leaves broadly elliptic to ovate, obtuse, imbricate, alternate, 2.5—5 mm 



77 



98 



long, entire; inflorescence few -flowered, consisting of 2 — 3 secund 
branches or spikes, with subsessile flowers; bracts subovate, 3—4 mm long; 
flowers 5-, less often 4-merous; sepals triangular, ca. 1 — 1.5 mm long, 
broader than long, glabrous, connate at base, acute; petals five, 2 — 3 times 
as long as calyx, lanceolate, 1 -nerved, whitish or pinkish, reddening along 
the nerve, free to base, 3—4 mm long, apiculate; stamens 5 (or 4), alternating 
with and shorter than petals, sometimes with 5 (4) more obsolete rudimentary 
stamens opposite to petals; hypogynous scales linear- spatulate, 0.5—0.75 mm 
long; follicles stellately spreading, connate at base, declinate, narrowly lanceo- 
late, longitudinally sulcate and ribbed, ca. 5 mmlong, many-seeded; seeds ovoid, 
obtuse, 0.6—0.9 mm long, 0.4—0.45 mm broad. April. (Plate V, Figure 4a). 

Sandy and stony soils on dry slopes.— European part: Crim. (Yalta, 
Alushta, Balaklava); Caucasus: Cisc. : Temyruk; Dag.: Makhachkala, 
Chir-yurt; W. Transc. : Black Sea shores; E. Transc. (Tbilisi, Baku), Tal. : 
Lenkoran. Gen. distr. : Bal. -As. Min., Med. Described from Montpellier. 
Type in London. 

Note. The reported finding ofS. rubrum (S.caespitosum)in 
the Kopet Dagh Range in Turkmenia relates to S. t e t r a m e r u m. 



Series 3. Macrosepalae A.Bor.— Stamens uniseriate, as many as petals; 
calyx exceeding corolla. 

53. S. tetramerum Trautv. in A. H. P. VII (1881 ) 454; Shmal'g., Fl. I, 362; 
Boiss., Fl. Or. Suppl. (1888)247.— Macrosepalum turkestanicum 
Rgl. et Schmalh., Izv. Obshch. lyubit. estestvozn. , antrop. i etnogr., XXXIV, 2 
(1882) 25; Fedtsch. O. et B., Consp. Fl. Turk. 3 (1909) 79.- Sedum 
turkestanicum Hamet in B. Fedch., Rast. Turk. (1915) 478 n. nud. — 
S. aetnense var. tetramerum Hamet in Bull. Jard. Bot. 1— 2 (l 914) 143.- 
Ic: Frod.in Acta Horti Gothoburg. VII (1932) textfig. 754-762. - Exs.: 
H. F. A. M., No. 426a, 426b. 

Annual; roots fibrous, slender; stems 2—5.5 cm long, glabrous, simple 
or slightly branching nearly from base, erect, slender, leafy from base; 
leaves alternate, semiterete, ovate, entire, 2.5—5 mm long, with a subquadrate 
appendage-spur at base, subacuminate, slightly tuberculate on the surface, 
glabrous or sparsely ciliate-margined; inflorescence 2— 8 -flowered, 
spiciform, Y 3 — x j 2 the length of the entire plant; bracts in pairs near each 
flower, ovate, ciliate, 3—4 mm long, about as large as leaves; flowers 
4-merous, subsessile, apparently axillary, crowded; sepals linear -terete 
or oblong-ovate, subobtuse, 4—6 mm long, 0.7—1 .5 mm broad, erect, with a 
basal spur, glabrous; petals ovate -lanceolate, 1 -nerved, acuminate, whitish, 
2— 3.5 mm long, shorter than calyx, tapering toward base and slightly fused; 
stamens 4, alternating with petals, more than half as long as and slightly 
adnate to petals at base, with rounded or reniform anthers; hypogynous 
scales linear-spatulate; follicles slightly divergent, linear-lanceolate, 
convex, subulate -pointed, as long as sepals, the surface densely glandular- 
tuberculate, scabrous, ca. 4 mm long; seeds ovoid, smooth, ca. 0.8 mm 
long, 4 mm broad. June— July. (Plate V, Figure 5a). 

Stony and pebbly soils in chalk mountains.— Caucasus: Dag. (Chir-yurt, 
Makhach-Kala), E. Transc. (Baku); Centr. Asia: Mtn. Turkm., T. Sh., 



78 



Syr.-D., Pam. -Al. Gen. distr. : probably in Iran. Described from the 
vicinity of Baku. Type in Leningrad. 

54. S. aetnense Tin. ap. Guss., Fl. Sic. synops. II, 2 (1844) 826; Berger 
in Engl. u. Pr., Nat. Pflzfm. 18a (1930) 460.- S. skorpili Velen. in 
Sitzber. Boehm. Ges.XL (1899), No. 2 9. - S. albenicum Beck in Ann. 
Nat. Hofmus.XIX (1904) 74.- S. erythrocarpum Pau in Bull. Ac. 
int. geogr. bot. (1908), No. 206.— S. aetnense var. genuinum Hamet 
in Bull. Jard. Bot., No. 1-2 (1914) 143.- Ic: Frod. in Acta Horti Gothoburg. 
VII (1932) pi. L, 6; textfig. 763-770. 

Annual; flowering stems, simple or branching from base, 2—5 cm 
high; leaves semiterete, lanceolate to oval, ciliate -margined or 
cartilaginous -dentate, 3—5 mm long; bracts oblong, ciliate, ca. 3.5 mm long; 
flowers 4 or 5 -merous, sessile, in leaf axils throughout the stem; sepals 
longer than petals, ciliate -margined, obtuse, subovate, ca. 4 mm long; petals 
obovate, somewhat narrowing toward base, short "acuminate, 2— 2.5 mm 
long, white; stamens 4, rarely 5, shorter than corolla, somewhat more 
than half as long as petals; hypogynous scales narrowly linear, 0.6X1.15 mm; 
follicles broad, connate at base, stellately spreading, convex ventrally, 
densely and minutely glandular -tuberculate on the surface, 3—3.5 mm long, 
many-seeded; seeds small, ca. 1 mm long, 0.4 mm broad, obtuse, oblong- 
ovoid. May. 

Sandy and stony soils.— European part: Bl. (near Nikolaev and Aleshka), 
Crim. (near Sudak), Alma River valley. Gen. distr.: Med., Bal.-As. Min. 
Described from Sicily (vicinity of Mt. Etna). Type in Naples. 



Genus 700. PSVJDOSEDUM (BOISS.) Berger 

Berger in Engl. u.Pr. Nat. Pflzfm. 18a (1930) 465.- Sect. Pseud osed um gen. u mb i li c us Boiss., Fl. Or. II, 

775(1872). 

Calyx divided nearly to base into 5 or 6 lobes; corolla pink or reddish, 
drying golden-yellow, connate nearly to the middle, infundibular or 
campanulate 5- or 6 -dentate; stamens 10—12; pistils 5 or 6, with long 
stules; fruit follicles, erect, lanceolate; seeds numerous, small, ca. 1 mm 
long, mostly oblong. Perennial, glabrous herbs, with caudex, very few small 
membranous triangular leaves, and cordlike or tuberous roots; fruiting 
stems densely leafy, erect or ascending at the base, simple, sometimes the 
old stems persistent; leaves alternate, fleshy, oblong to linear, terete; 
inflorescence umbellate -corymbiform, many -flowered. 

Species confined to mountainous habitats in T. Sh., Pam. -AL, Kopet Dagh, 
and the northern part of Iran; only one species grows in the foothill -plain 
part of the Central Asian semidesert and desert regions. 

1. Roots 1—4, tuberous, rounded or fusiform; corolla campanulate- 
infundibular, with oblong -lanceolate, declinate teeth erect in fruit. 

(T. Sh., Alai) 5. P. ferganense A. Bor. 

+ Roots clustered or solitary, not tuberous, slender or thickened 2. 

2. Root 1, vertical, thick or branching into several cordlike -thickened 
roots; corolla campanulate, with teeth half as long as tube ....... 3. 



79 



+ Roots numerous; corolla infundibular or narrowly campanulate, with 

teeth from longer to somewhat shorter than tube . . 5. 

3. Caudex 3— 4 cm long, with persistent old stems; calyx l / 2 — 2 / 3 as long 
as corolla, with linear -lanceolate, acute lobes (Kara-tau Range) . . . 
9. P. karatavicum A. Bor, 

+ Caudex short or obsolete; calyx %— / 3 as long as corolla, with 

oblong or oblong-lanceolate, obtuse sepals 4, 

4. Corolla 12 mm long; calyx Y 5 — % as long as corolla, with oblong- 
lanceolate sepals; all stamens shorter than corolla. (Kopet Dagh 
Range) 6. P. multicaule (Boiss. et Buhse) A. Bor, 

+ Corolla 8— 9 mm long; calyx */, as long as corolla, with oblong sepals; 

stamens unequal: 6 about as long as and 6 shorter than corolla 

. . 8. P. campanuliflorum A. Bor 

5. Densely cespitose plants; roots numerous, slender, with spherical or 
ovoid thickenings; sepals oblong-lanceolate, acute; corolla narrowly 

campanulate; stamens much shorter than corolla 

7. P. fedtschenkoanum A. Bor, 

+ Noncespitose plants; roots fasciculate; sepals oblong or ovate, 
subobtuse; corolla infundibular; stamens slightly shorter than or 
as long as corolla 6, 

6. Flowers 10— 14 mm long; corolla pink, drying golden-yellow, 
infundibular, with lanceolate teeth about as long as or slightly longer 
than tube 8, 

+ Flowers 7—10 mm long; corolla red or violet, often faded when dry, 
broadly infundibular, with lanceolate or oblong-lanceolate teeth 
slightly shorter than or as long as tube 7, 

7. Inflorescence corymbiform, broad, compact; corolla broadly 
campanulate -infundibular, with oblong-lanceolate teeth; stamens 

unequal: 6 as long as and 6 shorter than corolla 

3. P. condensatum A. Bor. 

+ Inflorescence corymbiform -paniculate, with many -flowered, divaricate 
branches; corolla infundibular, with lanceolate teeth; 6 stamens as 
long as and 6 longer than corolla 3. P. bucharicum A. Bor, 

8. Flowers 10—12 mm long; corolla teeth as long as or somewhat shorter 
1fl1 than tube. Plants 20—25 cm high; roots slender, to 2 mm in diameter, 

not tuberous (foothill plain of the northern part of Central Asia) 

1. P. lievenii (Ldb.) Berger, 

+ Flowers 12—14 mm long; corolla teeth surpassing tube. Plants to 

35 cm high; roots thick, to 7 mm in diameter, with few small tuberous 
thickenings 2. P. longidentatum A. Bor, 



Section 1 . P. LIEVENIA A. Bor. in Acta Inst. Bot. Ac. Sc. USSR. Ser. 1 , I 
(1933) 108.— Corolla infundibular or campanulate -infundibular; inflorescenc< 
branching, densely many -flowered; roots not tuberous. 

1. P. lievenii (Ldb.) Berger in Engl. u. Pr., Nat. Pflzfm. 18a (l 930 ) 465.- 
Umbilicus lievenii Ldb., Fl. Ross. II (1844-1846) 173, p. p. - 
U.steveni Ldb. in sched. - C o ty 1 e d o n lievenii Ldb., Fl. Alt. II (1830 ) 
57, 197.- Sedum lievenii Hamet in Candollea IV (1929-1 931 ) 38.- 



80 



S.inder iense Fisch. in sched. — S. virescens Schlechtend. in herb. — 
Ic: Ldb., Ic.pl. Fl. Ross. 1, 14 (1829) tab. 57; Frod. in Acta Horti Gothoburg. 
VI (1931) pi. XXVII, textfig. 308-316; A. Borissova in Acta Inst. Bot. Ac. Sc. 
USSR Ser. 1,1 (1933) 109, f. 1. 

Perennial; roots numerous, fascicular, slender, to 2 mm in diameter, 
more or less branching, tuberous thickenings absent; caudex robust; 
fruiting stems 1—4, 20—25 cm high; leaves linear, 0.5—2 cm long; 
inflorescence many -flowered, with numerous approximate branches 
elongating in fruit; lower branches arched-upcurved; bracts shorter than 
calyx, oblong or lanceolate, obtuse; pedicels 1—2 mm long; flowers 10— 12 mm 
long; calyx 3 mm long, x / 4 — J / 3 as long as corolla, divided nearly to base 
into oblong, subacute lobes; corolla indundibular, broadening in fruit, 
campanulate, partially rupturing, pink, drying golden-yellow, with 5 or 6 
lanceolate, acute, divergent teeth slightly shorter than or as long as tube; 
stamens 10— 12, the 5 or 6 opposite to sepals somewhat shorter than corolla, 
the 5 or 6 opposite to petals not exceeding middle of corolla teeth; anthers 
ovate, 0.9 mm long, sharp -pointed; hypogynous scales small, broad, with 
narrow base, cordate; pistils shorter than corolla, lanceolate, with 
subulate beak; seeds ellipsoid -oblong, tapering toward the apex, 1 mm 
long. April— May. 

Stony and clayey saline soils in semideserts and deserts.— W.Siberia: 
Irt.; Centr. Asia: Ar.-Casp., Balkh., Kyz. K. Gen. distr. : Dzu.-Kash 
(Kuldja). Described from Lake Inder. Type in Leningrad. 

Note. Specimens from the Nura-Tau and Mogol-Tau differ from the 
typical Pseudosedum lievenii, but owing to lack of material and the 
bad state of preservation, they cannot be separated. 

2. P. longidentatum A. Bor. in Acta Inst. Bot. Ac. Sc. URSS. Ser. 1 , I 
(1933) 109.- Ic: A. Bor., 1. c, f. 2. 

Perennial; roots numerous, to 7 mm in diameter, fasciculate, with 
small, sparse (remote) tuberous thickenings; caudex robust; fruiting 
stems 1—3, robust, erect, 25—40 cm high (rarely less tall); leaves linear, 
1—2 cm long, 1 mm broad; inflorescence 5—6 cm long, densely many- 
flowered, with numerous branches elongating in fruit; lower branches 
arched-upcurved; bracts oblong, obtuse, 2—4 mm long, shorter than calyx; 
flowers 12— 14 mm long; calyx 3— 4 mm long, Y 5 — % as long as corolla, 
divided nearly to base into oblong, subacute sepals; corolla infundibular, 
campanulate in fruit and partially rupturing, pink, drying golden-yellow, 
with 5 or 6 lanceolate, acute teeth surpassing corolla tube; stamens 10 — 12, 
the 5 or 6 opposite to sepals scarcely shorter than or as long as petals, the 5 or 6 
opposite to petals shorter than petals by Y 3 of length of corolla teeth; anthers 
ovate, 0.9 mm long, sharp -pointed; pistils shorter than corolla; seeds 
ellipsoid -oblong, attenuate toward the apex, 1 mm long. April— May. 

Alpine and subalpine meadows, slopes, among woody -shrubby vegetation, 
on steppe slopes, on pebbly soils, in mountains at (750 ) 1 ,500 — 3,000 m. — 
Centr. Asia: T. Sh., Pam.-Al. Endemic. Described from the vicinity of 
Alma-Ata. Type in Leningrad. 

3. P. condensatum A. Bor. in Acta Inst. Bot. Ac. Sc. URSS. Ser. 1, I (1933) 
110.- Ic: A. Bor., I.e., Ill, f. 3. 

Perennial; roots numerous, nearly fascicular, long, slender, 1—1.5 mm in 
diameter, often branching, with remote, small tuberous thickenings, 4 mm 



81 




PLATE VI. 1.— Pseudosedum ferganense A.Bor.: a) flower; 2 — P. mult icaule (Boiss.et Buhse) 
A.Bor.: a) flower; 3-Orostachys m a la co phylla (Pall.) Fisch.: a) flower; 4 — O. thy rsi fl o ra Fisch. 
a) flower, b) rosette leaf, c ) cauline leaf; 5-O.fimbriata (Turcz.) Berger: a) rosette leaf, b) cauline 
leaf. 



82 



.05 



broad, 5 mm long; caudex short or obsolete; fruiting stems 1—3, erect or 
ascending in lower part, 1.5—2 mm in diameter in middle part, 20— 25cm 
high; leaves linear, 1—2.5 cm long, 1 mm broad; inflorescence dense, 
compact, many-flowered; branches 1— 1.5 cm long, erect or slightly arcuate; 
calyx Y 4 — y 3 as long as corolla, 2— 3 mm long, divided nearly to base into 
ovate, subobtuse lobes; corolla campanulate and broadly infundibular, not 
inflating in fruit but rupturing into fragments, drying pale violet, with 6 
dark -violet, oblong-lanceolate, erect, acute, divergent teeth slightly shorter 
thantube; stamens 12, the 6 opposite to petals shorter than them by l j z the length 
of teeth, the 6 opposite to sepals as long as petals; anthers broadly ovate, 0.6 mm 
long, sharp -pointed; pistils about as long as corolla; seeds numerous, 
oblong, 1 mm long. May— July. 

Stony slopes, among shrubs, at ca. 2,000 m and to the alpine zone. — 
Centr.Asia: Pam.-Al. Endemic. Described from the W. Pamir (Shugnan). 
Type in Leningrad. 

4. P. bucharicum A. Bor. in Acta Inst. Bot. Ac. Sc. URSS, Ser. 1, I (1933) 
111.- Ic: A.Bor., I.e., f. 4. 

Perennial; roots numerous, nearly fascicular, slender, 1— 2 mm in 
diameter, branching, with sparse, small, oblong -terete, inconspicuous 
tuberous thickenings; caudex short; fruiting stems numerous, 2— 3 mm 
in diameter in middle part, erect, scarcely ascending, with persistent 
old stems, hollow, simple, 20 (30?) cm high; leaves linear, to 2 cm long, 2 mm 
broad, obtuse; inflorescence umbellate -paniculate, many -flowered, 
inflorescence branches 2— 3.5 cm long, spreading, the lower arched -upcurved, 
the others shorter and erect; bracts linear, obtuse, about as long as calyx; 
pedicels to 0.5 mm long; flowers 7— 10 mm long, subsessile; calyx % — l / 6 as 
long as corolla, divided nearly to base into oblong, obtuse, greenish lobes; 
corolla infundibular, campanulate in fruit, the 6 teeth declinate, lanceolate, 
acute, red, faded when dry, 4—5 mm long, as long as or slightly longer than 
tube; stamens 12, the 6 opposite to petals about as long as them, the 6 opposite to 
sepals as long as or slightly longer than petals; anthers dark violet, oblong, 1 mm 
long; styles half as long as corolla; pistils scarcely longer than corolla; 
seeds numerous, oblong, 0.5 mm long. April — May. 

Mountains, at 900-1,100 m. - Centr.Asia: Pam. -Al. (Mt. Khodzha-Mamyn, 
Kulyab, Bal'dzhuan). Endemic. Described from Tadzhikistan (Khodzha- 
Mamyn). Type in Leningrad. 



Section 2. TUBERARIA A. Bor. in Acta Inst. Bot. Ac. Sc. URSS, Ser. 1, I 
(1933) 112.— Roots tuberous; corolla campanulate -infundibular. 

5. P. ferganense A. Bor. in Acta Inst. Bot. Ac.Sc. URSS, Ser. 1, I (1933) 
112.- Ic: A. Bor., I.e., 113, f. 5. 

Perennial; roots 1—3, rarely 4, tuberous, rounded, 1 — 1.5 cm in diameter 
or fusiformly thickened (l— 1 .5 cm in diameter in upper part); caudex very 
short, simple or branching; fruiting stems 1—3 (5), slender, to 1.5 cm in 
diameter, erect or somewhat ascending, simple, 10—20 cm high; sterile 
stems densely leafy; leaves oblong, 8 mm long, 2 mm broad; inflorescence 
many -flowered, rarely 5— 7 -flowered, 1.5 cm long, 1.5—3 cm in diameter, 



83 



umbellate -corymbiform; inflorescence branches suberect or slightly 
angularly curved, crowded, 1—4 cm long, the lower mostly not arcuate; 
bracts lanceolate, acute, to 2 mm long, half as long as calyx; pedicels 
1—5 mm long; flowers 10 — 12 mm long; calyx Y 5 — / 4 as long as corolla, 
divided nearly to base into lanceolate, acute lobes; corolla campanulate- 
infundibular, campanulate in fruit, pink(?), drying pale pink, with 5 or 6 
oblong-lanceolate, subacute teeth intensely colored along the midnerve, 
declinate, in fruit slightly shorter than corolla tube; stamens 10— 12, the 5 or 6 
opposite to petals 2 / 3 their length, the other 5 or 6 opposite to sepals somewhat 
shorter than petals; anthers oblong, 1 mmlong, muticous; pistils as long as 
corolla; seeds oblong, tapering at the apex, 1 mm long. May — June. 
(Plate VI, Figure la). 

Dry terraces and stony slopes.— Centr. Asia: T. Sh. (Kirghizia), Pam. -Al. 
(Alai Range). Endemic. Described from the vicinity of Gulcha. Type in 
Leningrad. 



Section 3. CAMPANELLA A. Bor. in Acta Inst. Bot. Ac. URSS, Ser. 1, 
I (1933) 112.— Root not tuberous; corolla campanulate; inflorescence 
umbellate -corymbiform, little branching, twice as broad as long. 

6. P. multicaule (Boiss. et Buhse) A. Bor. in Acta Inst. Bot. Ac. Sc. 
URSS, Ser. 1, I (1933) 112.- Umbilicus multicaulis Boiss. et Buhse 
in Nouv. Mem. Soc. Nat. Mosc. XII (i860) 93.- Ic. : A. Bor., 1. c, 113, f. 6. 

Perennial; root l,more or less thickened, 5— 10 mm in diameter; caudex 
thickened, with persistent old stems; fruiting stems 7 or more, to 1.5 mm 
in diameter, erect, simple, 12—20 (25), mostly 15 cm high, densely leafy, 
1 — 1.5 mm in diameter; leaves fleshy, subterete, linear or linear-oblong, 
subobtuse, broadening toward base, 1—3 mm broad, 0.5 — 2 cm long; 
inflorescence umbellate -corymbiform, furcate, twice as broad as long, 
the branches divaricate, erect or slightly angularly curved, not elongating 
in fruit, 1.5—2 cm long, the lower branches not arcuate; bracts oblong, 
3—4 mm long, 1—1.5 mm broad, shorter than calyx; pedicels thickish, 
about as long as calyx; flowers 12 mm long; calyx Y 4 as long as corolla, 
green, divided nearly to base into obtuse, oblong sepals 3—5 mm long, 
1 mm broad; corolla campanulate, inflated in fruit, constricted below the 
teeth, pale red (pink) with 6 ovate, subacute teeth half as long as tube, 
slightly declinate, drying purple; stamens 12, shorter than corolla, the 6 
opposite to petals half as long as the teeth,the 6 alternate with petals, somewhat 
longer; anthers oblong, 1 mm long, sharp -pointed; pistils as long as 
corolla; seeds oblong, 1 mm long. April— June. (Plate VI, Figure 2a). 

Mountains, steppe zone, at 1,500—2,500 m. — Centr. Asia: Mtn. Turkm. 
(Kopet Dagh Range). Gen. distr. : Iran. Described from N. Iran. Type in 
Leningrad. 

7. P. fedtschenkoanum A. Bor. in Acta Inst. Bot. Ac. Sc. URSS, Ser. 1 , I 
(1933) 114.- Ic: A. Bor., I.e., 115, f. 7. 

Perennial; roots numerous, slender, tuberous, semispherical or oval, 
0.5—1 cm broad, 0.5—2 cm long. Densely cespitose plants with short 
caudex; stems numerous, slender, 1—2 mm in diameter, erect or somewhat 



84 



ascending, strong, hollow, simple, 7—20, mostly 12 cm high; old stems 
persistent; leaves linear, to 1 cm long, 1 mm broad, subobtuse; inflorescence 
umbellate -corymbiform, densely many -flowered, with branches 0.5—1 (l.5)cm 
long; lower branches somewhat arched -upcurved, the others erect; bracts 
oblong-lanceolate, acute, 3.5—4 mm long, 0.5 mm broad, shorter than calyx; 
pedicels thickish, scarcely 0.5 mm long or inconspicuous; flowers 7— 10 mm 
long, subsessile; calyx Y 3 as long as corolla, dry -brownish, divided nearly 
to base into 5 acute, oblong -lanceolate lobes, 3—4 mm long, slightly broadening 
toward base; corolla narrowly campanulate, with cylindrical tube and 5 
5 teeth, violet, drying pale violet or faded; corolla teeth more intensely 
colored, drying violet, 3—4 mm long, 2 mm broad at base, ovate, acute; 
corolla broadly campanulate in fruit, its teeth slightly declinate; stamens 10, 
shorter than corolla, the 5 opposite to petals attaining half the length of the teeth, 
the other 5 slightly longer; anthers oblong, dark violet, 0.9 mm long; 
pistils shorter than corolla; seeds numerous, oblong -pyriform. March- 
May. 

Stony mountain slopes at 1,000— 2,200 m. — Centr.Asia: Pam. -Al. 
(Kugitang, Gissar, Zeravshan, Bal'dzhuan, Kabadian, and Chalshau ranges). 
Endemic. Described from Tadzhikistan (Kafirnigan River). Type in 
Leningrad. 

8. P. campanuliflorum A. Bor. in Acta Inst. Bot. Ac. Sc. URSS, Ser. 1 , 
I (1933) 115.- Ic: A. Bor., 1. c, f. 8. 

Perennial; root 1, long, robust, 5—8 mm in diameter, vertical, with 
slender lateral roots; caudex obsolete; stems numerous, divergent, densely 
leafy, partly sterile, simple, 10— 15 cm high; leaves linear, 1—1.5 mm 
broad, subobtuse; inflorescence umbellate -corymbiform, 1.5 cm 
long, 2.5— 3.5 cm broad; branches short, 1.5 cm long, 2— 3 flowered, 
angularly curved, the lower not arcuate; bracts oblong-ovate, obtuse, 
shorter than calyx, to 1.5 mm long; pedicels to 1 mm long; flowers 8—9 mm 
long; calyx a / 3 as long as corolla, 3 mm long, divided nearly to base into 
oblong-lanceolate, obtuse lobes; corolla campanulate, pink, with petals 
more intensely colored along the midnerve, broadly campanulate in fruit, 
with 5 or 6 erect teeth half as long as or slightly longer than tube, to 3 mm 
long; stamens 10— 12, the 5 or 6 opposite to petals half the length of teeth,the 
other 5 or 6 about as long as corolla; anthers ovate or broadly ovate, 
0.9 mm long; pistils shorter than corolla; fruit as long as corolla; seeds 
numerous, oblong -ovoid, 1 mm long. April— May. 

Stony soils on mountain rocks.— Centr.Asia: Pam. -Al., Dzhizak, 
Ingyrchak Pass). Type in Leningrad. 

9. P. karatavicum A. Bor. sp. nova in Not. syst. VII, 8 (1938) 185. 
Perennial; root long, robust, branching into a series of cordlike thickened 

long lateral roots, ca. 0.4—0.5 cm in diameter; caudex branching, 3—4 cm 
long, enveloped by numerous old stems and by membranous, broadly 
triangular leaves ca. 3 mm long; stems numerous, slightly divergent, densely 
leafy, partly sterile, simple, 5—10 cm high; leaves linear, ca. 0.5 cm long, less 
than 1 mm broad, somewhat narrowed toward the apex; inflorescence 
umbellate -corymbiform, compressed, few -flowered, ca. 1.5 cm in diameter, 
1 cm long, with short branches; bracts lanceolate, subobtuse, 0.5 cm long, 
ca. 1—1.5 mm broad; flowers ca. 1 cm long, with very short pedicels or 



85 



subsessile; calyx 1 / 2 — % as long as corolla, divided nearly to base into 
linear-lanceolate, acute lobes; corolla campanulate, pink, drying pinkish 
or yellow, with 5 straight, broadly triangular teeth 1.5 times as long as 
tube; stamens 10, those opposite to petals half the length of corolla, the other 
5 about as long as corolla; anthers oblong, slightly more than 1 mm long; 
pistils shorter than corolla, with subulate erect style; follicles lanceolate, 
shorter than corolla, with long, slender beak; seeds oblong, less than 1 mm 
long, numerous. June. 

Stony and pebbly soils. - Centr. Asia: T. Sh. (Kara-Tau Range). Endemic. 
Described from the Kara-Tau Range. Type in Leningrad. 

Note. Closely related to P. c a m p a n u 1 i f 1 o r u m A. Bor., it is 
distinguished by its long caudex covered with persistent old stems, leaf 
size, different calyx and corolla, and other characters. 



Genus 701. OROSTACHYS (DC.) FISCH. 

Fisch.Cat. Gor. (1808) 99; Berger in Engl. u.Pr. Nat. Pfzfm. 18a (1930) 463.- Genus Umbilicus sect.oro- 
stachys DC.Prodr. Ill (1828) 400; Ldb., Fl. Ross. II (1844-1846) 173.- Genus Cotyledon sect. Or os t a c hy s 
Schoenl.in Engl.und Prantl Nat.Pflzfm.l Aufl.III, 2a (1890) 33. 

Flowers 5 -merous, yellowish, greenish, white, pinkish, or red; sepals 
fleshy, V 3 — ! / 2 as l° n g as corolla; petals free or short -connate at base, 
triangular-lanceolate, acute; stamens 10, filiform, with elongate-rounded 
anthers; hypogynous scales small, truncate at the tip; follicles slightly 
inflated, free, broadening only at base, with a slender, long beak at the 
apex; seeds numerous, linear. Glabrous plants, 5—30 cm high, seldom 
higher; in first year leaves in basal rosettes, usually cuspidate from a 
white cartilaginous apical appendage, rarely soft -acuminate or subobtuse, 
linear to ovate; in the second year a solitary flowering shoot arises 
from center of rosette; flowers subsessile or with more or less 
conspicuous pedicels, numerous, in dense pyramidal racemes or paniculate 
inflorescences. 

Species distributed from the Urals to China and Japan, in Central Asia 
as far south as Tien Shan and the Pamir-Alai. 

1. All leaves muticous, elliptic or ovate to oblong and oblong-lanceolate, 
subobtuse or soft -acuminate; flowers yellow or greenish white; 
bracts broad, ovate. (Section 1 . Euappendiculata A. Bor.) 
1. O. malacophylla (Pall.) Fisch. 

+ Cauline leaves cuspidate; rosette leaves cuspidate from a white 
cartilaginous appendage; bracts oblong to linear. (Section 2. 
Ap p e nd i c u 1 a t a A. Bor.) 2. 

2. Cartilaginous appendage at tip of rosette leaves entire, sometimes 
inconspicuously sinuate (Plate VI, Figure 4b); flowers greenish, 
yellowish, white, pinkish, or with reddish-tipped petals 3. 

+ Cartilaginous appendage at tip of rosette leaves spiny -dentate 

(Plate VI, Figure 5a); flowers reddish, drying blue 

5. O. fimbriata (Turcz.) Berger. 

3. Flowers white or greenish white, pinkish or with petals reddish at 
tip and in buds, conspicuously pedicellate, less often subsessile, often 
several on each pedicel; stamens with dark anthers 4. 

* From the Greek oros, mountain and st a chys, spike. 

86 



+ Flowers greenish yellow, each one with a very short pedicel or sessile; 

inflorescence compact, dense; stamens with yellow anthers 

2. O. spinosa (L.) C. A. M. 

4. Flowers white or white-pink, with stamens as long as or longer than 

corolla; leaves light green; bracts shorter than flowers, ovate -oblong 

(SE of the RSFSR, W. Siberia, NE Central Asia) 

3. O. thyrsiflora Firsch. 

+ Flowers greenish white or reddish, often macular; stamens shorter 

than corolla; leaves dark green, often reddish, glaucous, macular; 

bracts as long as or longer than flowers, lanceolate or linear. 

(Far East) 4. O. cartilaginea A. Bor. 



Series 1. Euappendiculatae A. Bor.— Leaves unarmed, subobtuse or 
soft -acuminate, all without cartilaginous appendage. 

1. O. malacophylla (Pall.) Fisch. in Mem. Soc. Nat. Mosc. E (l 809) 274; 
Berger in Engl. u. Pr., Nat. Pflzfm. 18a (1930) 464. - O. s errata Sweet 
Hort.Brit. ed. II (1 830 ) 22 5.- Cotyledon malacophyllum Pall., It. Ill 
(1776) 266, 320 app. 729, No. 88; Kom. and Alis., Opr. Dal'nevost kraya I, 
601.- Umbilicus m ala c ophy 1 1 u s DC, Prodr. Ill (l 828) 400; Turcz., 
Fl. baic.-dah. 1,433; Ldb., Fl. Ross. II, 174; Maxim., Pr. Fl. Amur. (1859) 
114.- U. st amine us Ldb., Fl. Ross. II (1834-1846) 174.- Sedum 
malacophyllum Steud., Nomencl. (l 821 et 1841) 759; Franchet, PL 
Davidian. I (1884) 129; Frod. in Acta Horti Gothoburg. VI (l 931 ) 9 - 
Ic: Kom. and Alis., 1. c, tab. 180, 1-3; Pallas, 1. c, tab. O, f. 1. Frod., 1. c, 
pi. I— III, textfig. 1-8. 

Biennial, in the first year with rounded rosettes of crowded leaves; 
rosette leaves without apical spiny prickle, obtuse or shortly soft- 
acuminate, oblong-lanceolate, obovate, elongate -elliptic or elliptic, entire; 
in the second year solitary stems (5) 10— 30 (35) cm long arise from 
center of rosette: cauline leaves alternate, approximate, overlapping, 
larger than rosette leaves, to 7 mm long, soft -acuminate; inflorescence 
a dense, spiciform, elongated raceme, sometimes branching; bracts 
broad, spatulate -ovate, often dentate, short -acuminate in upper part, 
mostly covering the flowers; flowers subsessile, numerous; calyx with 
acute, oblong sepals, 3—4 mm long, connate at base; petals greenish 
white or yellowish, connate at base, oblong or ovate, often crenate at the 
tip, acute, 4—6 mm long; stamens slightly or much longer than corolla, 
with yellow anthers; hypogynous scales subquadrate, obtuse or 
emarginate; follicles many-seeded, broad, ovate, tapering toward base 
and toward apex, about as long as corolla, with subulate beak 1 / 5 — / 6 as 
long as fruit; seeds ovoid, small. Fl. from end of August — September. 
(Plate VI, Figure 3a). 

Sandy, pebbly seashore embankments and riverbanks, rocks, dry 
gravelly taluses, rock crevices; nearly always occurring in large groups. — 
E.Siberia: Lena -Kol. , Ang. -Say., Dau.; Far East : Uda, Ze.-Bu., Sakh. 
Gen.distr.: Mong., Jap. -Ch. Described from Dauria. Type in London (?). 



87 



Series 2. Appendiculatae A.Bor.— All leaves spiny-acuminate; rosette 
leaves with cartilaginous appendage, 'entire or spiny-dentate, cuspidate. 

2. O. spinosa (L.) C. A. M. in Ldb., Reise (1830) 496; Berger in Engl, 
u. Pr., Nat. Pflzfm. 18a (1930) 464. - O. spinosa Sweet, Hort. Brit. ed. II 
(1830) 225.- O.chlorantha Fisch. in Mem. Soc. Nat. Mosc. II (1809) 274.- 
Cotyledon spinosa L., Sp. pi. (1753) 429; Ldb., Fl. Alt. II, 200; Kom. 
and Alis., Opred. rast., Dal'nevost. kraya I, 601 ; Fedtsch., Consp. Fl. Turk. 

3 (1909) 70; Turcz. in Bull. Soc. nat. Mosc. I (1938) 92. - C r a s s ula 
spinosa L., Mant. II (l 771 ) 388; Willd., Sp. pi. II, 1534. - U mb i 1 i c u s 
spinosus DC, Prodr. Ill (1828) 400; Ldb., Fl. Ross. II, 174; Maxim., 
Pr. Fl. Amur. (1859) 114; Kamch., Fl. Zap. Sib. VI, 1402; Turcz., Fl. 
baic.-dah. I (1842-1845) 432.- U. (Orostachys) erubescens Maxim., 
Prim., fl. Amur. (1859) 114.- Sedum spinosum Thunb., Fl. jap. (1784) 
186.— S. spinos um Willd., Enum. hort. Berol. (l 809) 485; Frod. in Acta 
Horti Gothoburg. VI (l 931 ) 13. — Semp ervivum cuspidatum Haworth, 
Miscell. Naturalia (1803) 186.- Ic: 1. c, pi. V, textfig. 41-59. 

Biennial, in the first year with rounded rosettes of crowded leaves, 
2—7 cm in diameter, with contracted stem, compactly enveloped by crowded 
leaves; rosette leaves fleshy, oblong, angularly rounded at the apex, with 
a whitish cartilaginous margin, abruptly passing into a cartilaginous 
prickle 2—4 mm long; in the second year a stem (rarely also lateral stems) 
arises from center of rosette, usually not branching, 10—30 cm high, with 
alternate, sessile, lanceolate leaves 1—2.5 cm long, 2—5 cm broad, gradually 
acuminate, with a cartilaginous prickle; inflorescence at end of stem — 
a long, many -flowered, compact raceme 5—20 (28) cm long; pedicels to 1 mm 
long or flowers sessile, with lanceolate or oblong bracts; calyx ca. 3 mm 
long, deeply divided into 5 lanceolate, acute lobes; corolla 2—3 times as 
long as calyx, greenish yellow, with ovate -lanceolate, acuminate petals, 
connate at base to % — 1 / 3 ; stamens longer than corolla, with yellow anthers; 
hypo gy nous scales short, subquadrate, slightly emarginate; follicles 
erect, glabrous, with sparse short hairs confined to the inner suture, 
lanceolate, 5—6 mm long, ca. 1 mm broad, with straight subulate beak about 
/ 3 as long as fruit; seeds oblong-ovoid, ca. 1 mm long, 0.2 mm broad. 
July— September. 

Open stony and gravelly slopes and rocks, in rock crevices in the 
steppe zone; often on solonetz and sandy soils in steppe plains; in southern 
open pine forsts and at their edges; in the forest zone only in southern 
part, confined to open southern slopes.— European part: V. -Kama, 
(S.Urals); W.Siberia: U. -Tob., Irt., Alt.; E.Siberia: Yenis., Ang. -Say., 
Dau.; Far East: Okh., Uda, Ze.-Bu., Uss.; Centr.Asia: T. Sh., Balkh., 
Dzu. -Tarb. Gen. distr. : Mong., Jap. -Ch., Dzu. -Kash., Tib. (W. part). 
Described from Siberia. Type in London. 

Note. Ledebour in Fl. Ross. II, 173, cites var. polystachys — 
a branching form of O. s p i n o s a. 

3. O. thyrsiflora Fisch. in Cat. Hort. Gorenk. (1808) 33; Mem. Soc. Nat. 
Mosc. II (1809) 274; Berger in Engl. u. Pr., Nat. Pflzfm. 18a (1930) 464.- 
O.roseus Berger in Engl., 1. c. — U mb i 1 i c u s thyrsiflorus DC., 
Prodr. Ill (1828) 400; Ldb., Fl. Ross. II, 174.- U. leucanthus Ldb., 1. c, 
173 (1844-1846); Kryl., Fl. Zap. Sib. VI, 1403.- Cotyledon thyrsiflora 



Maxim, in Bull. Acad. Petersb. XXIX (1884) 123.- C.leucantha Ldb., 

Fl. Alt. II (1830) 198.- C. rosea Less, ex Linnaea IX (1834) 177. — 

C. s errata Pall., Reise I (l 771 ) 256 (?) ex Fisch. (non L. ?). - C. spino sa 

Clarke in Hook., Fl. Brit. India II (1897) 416, non L. — Sedum spinosum 

var. thyrsiflorum Frod. in Acta Horti Gothoburg. VI (l 931 ) 15. — 

Ic: Ldb., Ic.pl. Fl. Ross. IV, tab. 395 (sub C. leuc antha); A. Borisova 

in Fl. Yugo-Vost. V, 475; Frod., 1. c, pi. IV, textfig. 60-66. 

Biennial; in the first year (before anthesis) with rounded rosettes of 
crowded leaves; rosette leaves light green, fleshy, oblong -triangular, 
concave, abruptly passing into a cartilaginous cuspidate appendage, 
1.5—2 mm long, imbricate; during anthesis an erect stem, 5—20 cm high, 
arises from center of rosettes, the stem covered with more or less remote 
leaves keeled on the back, hence subtriquetrous, oblong (except when dry), 
gradually passing into a cusp to 1— 1.5 mm long; inflorescence a long, 
many -flowered raceme; flowers several on each more or less elongated 
pedicel, pedicels of lower flowers 3—8 mm long, those of upper flowers 
smaller; bracts ovate -oblong, acuminate, shorter than flowers; calyx / 3 
as long as petals, with acute sepals, connate to the middle; corolla white 
or white-pink, connate at base, with 5 oblong petals 5—6 mm long; stamens 
10, with dark purple anthers, of which 5 longer than or about as long as 
corolla, the 5 opposite to petals shorter than corolla; filaments adnateto corolla; 
follicles oblong, gradually passing into beak, smooth, erect, as long as 
corolla; seeds small, ovoid. July — August. (Plate VI, Figure 4a— c). 
Rocks, stony slopes, solonetz soils, pebbly sites, desert steppes, sometimes 
sandy soils.— European part: Transv. (single — in Ilek District, in Chkalov 
[Orenburg] Region, near Bakaika and Burenino villages), V.-Kama (vicinity 
of Sterlitamak Lake, Asli-kul'); W. Siberia: U. Tob. (Chelyabinsk District, 
upper course of the Ural River, Guberlya Mts.), Alt.; Centr. Asia: widely 
distributed in Ar.-Casp., Balkh., Dzu.-Tarb., T. Sh., single-in Pam.-Al. 
Gen. distr. : Dzu. -Kash., Mong., Tib. Described from the Guberlya 
Mountains. Type in Leningrad. 

4. O. cartilaginea A. Bor. sp. nova in Addenda VIII, p. 3 68.— Cotyledon 
j apo nic a Kom. in Fl. Manch. II (1904) 402, p. p. (non Maxim). 

Biennial, with basal leaf rosettes in first year; rosette leaves oblong- 
lanceolate, flat, with white cartilaginous, convex, entire or scarcely crenate 
apical appendage, ca. 2—3 mm broad, 2 mm long, abruptly passing into a 
white cartilaginous prickle ca. 2 mm long; in the second year flowering 
shoots, usually not branching, arise from rosette, 10—20 cm high, with 
alternate, sessile, linear or lance-linear leaves, 1.5— 3.5cm long, 2—4 mm 
broad, gradually acuminate, with a white, cartilaginous filiform prickle; 
all leaves dark green or purple-dotted; inflorescence a terete, many- 
flowered, compact raceme, 3—15 cm long, sometimes branching; pedicels 
shorter than calyx or flowers subsessile; bracts linear or linear -lanceolate, 
about as long as or longer than flowers, mucronulate, usually macular; 
calyx ca. 2 mm long, macular, green, cut nearly to base into narrowly 
lanceolate acute lobes, sometimes mucronulate; corolla 5 -merous, white, 
greenish white or pinkish, reddish-tipped when in bud, often macular, 
3 times as long as calyx; petals connate to l / 3 , oblong-lanceolate, subacute; 
stamens 10, nearly all of equal length, slightly shorter than corolla, with 



89 



brown, later darkening anthers; follicles lanceolate, short -pediceled, with 
ca. 2 mm long filiform beak; seeds small, ca. 0.5 mm long, ovoid, brownish, 
numerous. Fl. August — September, fr. September. 

Rocks, taluses, ledges. — Far East: Uss. (Poset area, Novokievskoe 
village, Suifun River near Faddeevka village, Khunchun frontier post). — 
Gen. distr. : Jap.-Ch. (Korea, Manchuria, Liaotung Peninsula near 
Alsankiang railroad station). Described from the Suifun River, near 
Faddeevka village. Type in Leningrad. 

Note. This species has been confused with O. j a p o n i c a (Maxim) 
Berger, from which it is clearly distinguished by the cartilaginous appendage 
of the rosette leaves. In O. japonic a all leaves have a sharp subulate 
point, but without cartilaginous appendage — distributed in Japan, Korea, and 
Manchuria, but unknown in the Soviet Union. O. cartilaginea A. Bor. is 
closely related to O. m i n u t a Kom., described from Korea, but is 
differentiated by its larger size, shape of the cartilaginous appendage, 
larger bracts, and other characters. 

5. O.fimbriata (Turcz.) Berger in Engl. u. Pr., Nat. Pflzfm. 18a (1930) 
464.- Cotyledon fimbriata Turcz., Cat. pi. , baic. -dahur. (1838), 
No. 469; C.serrata L. (?), Sp. pi. (1753) 429. - U mb i 1 i c u s fimbriatus 
Turcz., Fl. baic. -dahur. I (1842-1845) 432.- U. denticulatus Turcz. in 
sched. — U. r a m o s i s s i m u s Maxim., Fl. Amur, (l 859) 472 in adnot. — 
Sedum fimbriatum Franchet in Nouv. Arch. Mus. Paris VI (1883—1884) 
12 8; Frod. in Acta Horti Gothoburg. VI (l93l) 11.— S.ramosissimum 
Franchet; 1. c, 128.- Ic: Frod., 1. c, textfigs. 23-32. 
1 14 Biennial, with rosettes of crowded leaves during first year of flowering; 

rosette leaves oblong, with white spines, broadening at base into white, 
cartilaginous, broad, spiny -dentate, lunate appendage; in the second year 
during anthesis, stem 10— 15 cm high, the leaves alternate, remote, with spines 
but not dentate, linear or lanceolate, 1— 3 cm long; inflorescence a dense 
raceme often branching from base; bracts spiny -acuminate; flowers 
usually long-pediceled, shorter -petioled in upper part of inflorescence; 
calyx with acuminate, oblong, green lobes 1—3 mm long; petals connate 
at base, reddish, lanceolate, entire, short -acuminate, 5—6 mm long; stamens 
usually shorter than or as long as petals; hypogynous scales quadrate or 
broadly spatulate; follicles oblong, with slender beak : / 4 as long as fruit; 
seeds small, ovoid, numerous. August. (Plate VI, Figure 5a— b). 

Stony soils in steppes.— E.Siberia: Dau. (Nerchinskii Zavod, Aga Steppe, 
Onon River basin). Gen.distr.: Tib., Mong., Jap. -Ch. Described from 
Dauria. Type in Leningrad. 

Genus 702. ROSULARIA * (DC.) STAPF 

Stapf in Bot. Mag. (1923) sub tab. 8985; Berger in Engl. u.Pr. Nat. Pflzfm. 18a (1930) 465.- U m b i li cu s sect, 
rosularia DC. Prodr.III (1828) 399p.p. 

Flowers 5—6 (9)-merous, whitish, pinkish, red, yellow; sepals fleshy, 
connate at base; corolla more or less high-connate, campanulate, the petals 
erect, acute, keeled on the back; stamens 10 — 12 (18), united with petals 
for much of their length; hypogynous scales more or less quadrate; 

* From the Latin rosul a, rosette — basal leaves rosulate. 



90 



follicles free, erect, gradually passing into beak. Low plants, glabrous 
or pubescent, with basal leaf rosette; leaves flat, sessile; flowering stems 
solitary when arising in center of rosette, numerous when arising 
laterally from axils of rosette leaves; inflorescence corymbiform- 
paniculate, paniculate, or racemose -paniculate, loose, sometimes the 
flowers solitary. Herbs resembling S e mp e r vi v u m, tuberiferous. 
Species distributed mainly in Central Asia and the Caucasus, as far east 
as Altai and as far west as Asia Minor, inclusive. 

Economic importance. Nearly all species of this genus are excellent 
as ornamental plants in carpetlike flower beds. 

1. Annuals or biennials, with slender, cordlike roots; in first year only 

few rosettes of basal leaves present; in second year solitary flowering 
shoots arise from center of rosette; inflorescence corymbiform; 
stems densely leafy. (Caucasus, Section Sempervivoides 

Boiss.) 2. 

Perennials, less often biennials, with tuberous or fusiform under- 
ground parts; inflorescence corymbiform, umbellate, paniculate, or 

racemose; stems with remote leaves 3. 

Petals dark red, lanceolate 2—2.5 times as long as red calyx; 
flowering shoots (5) 10—26 cm long, fleshy, thick, densely covered 

with ovate -cuneate leaves 10—30 mm long, 8—13 mm broad 

2. R. sempervivoides (Fisch.) A. Bor. 

+ Petals pinkish, pink -violet or whitish, 1.5 times as long as green calyx; 
flowering shoots 5—8 cm high, slender, covered with lanceolate -oblong 
leaves 7—10 mm long, 3—5 mm broad 1 . R. pilosa (M. B.) A. Bor. 

3. Inflorescence long, racemose or paniculate, dense, many -flowered; 
flowers small, 3—5 (6) mm long; corolla broadly campanulate, usually 
yellow, less often red. (Central Asia. Section O r i e nt a 1 e s A. Bor.) 
5. 

+ Plants not as above 4. 

4. Underground parts tuberous, mostly rounded; inflorescence corymbi- 
form or umbellate, loose; corolla broadly campanulate with broad 
base (3)6 — 10 mm long, white, yellow -violet, pinkish, violet, or yellow 
but then flowers small and plant 3—5 cm high; rosette leaves mostly 
ovate or oblong, obtuse or acute. Plants glabrous or puberulent 

or the leaves only ciliate -margined. Central Asia. (Section 

Campanella A. Bor.) 8. 

+ Underground part thickened, fusiform; inflorescence paniculate, loose; 
corolla infundibular -campanulate or campanulate, mostly with cuneate, 
narrow base, 5—10 mm long, yellow, red, pink; rosette leaves oblong 
or lanceolate, spatulate, obtuse. Plants mostly glandular -pubescent, 
less often glabrous. (Mainly Caucasian species, one species in Central 
Asia. Section Eu-rosularia Berger) 13. 

5. Inflorescence a long, narrow, racemose panicle or else narrowly 
paniculate, 2—3 (4) cm in diameter 6. 

+ Inflorescence broadly paniculate, 4—7 cm broad 7. 

6. Glabrous plants; calyx 1 / 3 as long as corolla; petals lanceolate; 

pedicels about as long as flowers, ca. 5— 6 mm long 

9. R. paniculata (Rgl. et Schmalh.) Berger. 



91 



116 



+ Glandular-pubescent plants; calyx half as long as corolla; petals 

oblong-elliptic; pedicels ca. 1— 1.5 mm long 

10. R. subspicata (Fr. et Sint.) A. Bor. 

7. Entire plants glabrous; corolla 5—8 mm long, yellow or yellow-green; 
sepals ovate 11. R. glabra (Rgl. et Winkler) Berger. 

+ Leaves hispidulous -ciliate, later glabrous; corolla ca. 4 mm long, 
purple, fleshy-red or whitish with purple glands, drying yellowish, 

with violet petal tips; sepals lanceolate 

12. R. turkestanica (Rgl. et Schmalh.) Berger. 

8. Corolla yellow. Plants 3—5 cm high 9. 

+ Corolla white, violet or yellow -violet (drying lilac in the middle, yellow 

at the edges). Plants (5)8-15 (l8)cm high 10. 

9. Flowers small, ca. 3 mm long, short -pediceled, in few -flowered 
inflorescence; petals narrowly lanceolate; cauline leaves acuminate 
13. R. kokanica (Rgl. et Schmalh.) A. Bor. 

+ Flowers 5 — 7 mm long, long -pediceled, in loose, many-flowered, 
corymbiform -umbellate inflorescence; petals oblong-lanceolate; 
cauline leaves subobtuse 14. R. lutea A. Bor. 

10. Corolla white (often drying pale yellow), connate into tube as long as 
calyx; rosette leaves rhomboid -ovate, subobtuse or concave at the 
apex, puberulent 18. R. platyphylla (Schrenk) Berger. 

+ Plants not as above ' 11. 

11. Petals 5— 7 (8) mm long, yellow -violet (drying dark violet); stamens 
with yellow filaments and anthers, later darkening; rosette leaves — 
mainly young leaves — white, long -ciliate, hispid on the margin, later 

glabrescent -margined, glabrous on the surface 

17. R. tadzhikistana A. Bor. 

+ Petals 6—10 mm long, white, pinkish or violet, darkening along the keel; 
stamens with white filaments and dark anthers 12. 

12. Rosette leaves ovate -oblong and — like the cauline — hispidulous on 
margin and surface; flowers mostly 5-, more rarely 6- or 7-merous; 
calyx Y 3 as long as corolla, with long white hairs at base; follicles 
pubescent 16. R. schischkinii A. Bor. 

+ Rosette leaves oblong, glabrous like the cauline, hispidulous -margined 
only at the summit; flowers mostly 6-, less often 7- or 8-merous; 

calyx half as long as corolla, glabrous; follicles glabrous 

16. R.alpestris (Kar. et Kir.) A. Bor. 

13. Plants more or less glandular -pubescent; corolla pubescent on the 
outside 15. 

+ Plants glabrous; corolla glabrous 14. 

14. Corolla yellow, ca. 7 mm long, with lanceolate -triangular petals; 
calyx 2— 2.5 mm long; fruit ca. 6 mm long, linear 

11? 7. R. lipskyi A. Bor. 

+ Corolla pink, ca. 10 mm long; sepals ca. 4-5 mm long; follicles 

ca. 9 mm long, elongate -lanceolate 5. R. persica (Boiss.) Berger. 

15. Flowering stems arising from axils of central leaves of radical rosette, 
mostly branching from base, forming a broad, many -flowered, 
thyrsoid inflorescence, with secund branches and with flowers from 

the very base; corolla red, ca. 5 mm long, connate nearly to the 

middle 6. R. elymaitica (Boiss. et Hausskn.) Berger. 



92 



+ Flowering stems arising in axils of lateral leaves of rosette; 

inflorescence corymbiform -paniculate, terminal; corolla 6 — 10 mm 

long 16. 

16. Calyx 2 / 3 as long as corolla, with broad base; corolla 6 mm long, 
broadly campanulate. Entire plant densely spreadingly glandular - 
pubescent. (Central Asia) 8. R. hissarica A. Bor. 

+ Calyx 2 / 5 — Y 3 as long as corolla, cuneate at base; corolla ca. 10 mm 

long. (Caucasus) 17. 

17. Flowering stems glandular -pubescent in upper part, glabrous in lower 
part; corolla whitish with dark glands 4. R. radiciflora (Steud.) A. Bor. 

+ Entire plant minutely villous -glandular; corolla reddish 

3. R. sempervivum (M. B.) Berger. 



Section 1. SEMPERVIVOIDES Boiss., Fl. Or. II (1872) 776.- Sect. 
Prometheum Berger (sub S e d o) in Engl. u. Pr., Nat. Pflzfm. 18a (l 930) 
459.— Annuals or biennials with cordlike, nonthickened roots; inflorescence 
corymbiform; flowers 5-merous; monocarpic; stems densely leafy. 
Caucasus. 

1. R. pilosa (M. B.) A. Bor. comb. nova. — S ed u m pilosum M. B., 
Fl.taur.-cauc. I (1808) 352; Boiss., Fl. Or. II, 786; Berger in Engl. u. Pr., 
Nat. Pflzfm. 18a (1930) 459; Gross., Fl. cauc. II, 459; Hamet in Acta Horti 
Tiflis. VIII, 3 (1908) 28.- Cotyledon pubescens C. A. M., Enum. pi. 
Cauc. (1831) 150.- Umbilicus pubescens Ldb., Fl. Ross. II (l 844-1 846) 
175.— Sedum regelii Hort. — Ic. : Bot. Mag. tab. 8503; Praeger in 
Journ. Hort.Soc. (l92l) f. 166; Frod. in Acta Horti Gothoburg. VII (1932) 
pi. IX, textfig. 64-71. 

Biennial; roots slender, fasciculate-cordlike; first year stems not 
developed in the form of a rosette of radical leaves; fruiting second -year 
stems 5—8 cm high, leafy, straight or slightly curved, mostly simple, 
glandular -pubescent; rosette leaves ovate -spatulate, the cauline lanceolate- 
oblong, all leaves glandular-pubescent, flat, fleshy, sessile, 7—10 mm long, 
3— 5 mm broad, obtuse, bright green; inflorescence umbellate -corymbiform, 
11fi 6 — 18 -flowered, dense, branching, 2— 3 cm long, 2— 4 cm broad; flowers 

5-merous; pedicels 4— 6 mm long; sepals connate at base, linear-lanceolate, 
acuminate, ca. 3.5—5 mm long, 1 — 1 .5 mm broad, glandular-pubescent, erect, 
green; corolla glabrous; petals pinkish or pink -violet or whitish, 1.5 times 
as long as calyx, 6—8 mm long, 2—3 mm broad, oblong-ovate, subacute, connate 
at base; stamens 10, slightly more than half as long as petals, with yellowish- 
reddish anthers; hypogynous scales less than 1 mm long, oblong -ovate, 
emarginate, colorless; follicles erect, green, as long as stamens, gibbous, 
divergent on the inner side, with reddish beak; seeds oblong, striped, 
ca. 0.8 mm long, ca. 0.3 mm broad. May— June. 

Rocky sites at 1,000— 2,300 m. Caucasus: Cisc. (northern slope of 
Main Range), Dag., E. Transc. (often), W. Transc. (rarely), S. Transc. (often), 
W. Transc. (rarely), S. Transc, Tal. (Zuvant). Gen. distr. : Arm. -Kurd, 
(former Kars Region, former Artvin District), Bal. -As. Min. Described 
from the Central Caucasus. Type in Leningrad. 

Note. Ornamental species suitable for carpetlike flower beds. The 
indication of S. pilosum for the Kopet Dagh Range near Kizyl-arvat is 



93 



erroneous (O. Kuntze in A. H. P. X (1887-1889) 189; Fedtsch. O. et B., Consp. 
Fl. Turk. 3 (1909) 73). 

2 . R. sempervivoides (Fisch. ) A. Bor . comb. nova. — Sedum semper- 
vivoides Fisch. ex M. B., Fl. taur. -cauc. Ill (181 9) 313; Hamet in Acta 
Horti Tiflis. VIII, 3 (1908) 26; Berger in Engl. u. Pr., Nat. Pflzfm. 18a (1930) 
459; Grossh., Fl.Cauc II, 229.- S.sempervivum DC, Prodr. Ill (1828); 
404; Ldb., Fl. Ross.II 185; Boiss., Fl. Or. II, 786. - S. d i v a r i c a t u m 
Schlecht. in herb. Berol. ex Ldb., Fl. Ross. II, 185; Boiss., Fl. Or. II, 786. - 
S. divaricatum Schlecht. in herb. Berol. ex Ldb., Fl. Ross., 1. c. — 
Ic: Gartenfl.t. 551, 1155; Bot. Mag. tab. 2174; Praeger in Journ. Hort. 
Soc. (1921) f. 165, Frod.in Acta Horti Gothoburg. VII (1932) pi. IX, 3-4; 
textfig. 74-81 . - Exs. : Herb. Fl. cauc, No. 323. 

Annual or biennial; roots cordlike, arising in a bundle from the tuberous 
underground part; first year stems short, arising from basal rosette, 
ca. 3—5 cm in diameter; the second -year flowering shoots arise, 10—26 cm 
long, erect, robust, densely leafy, pubescent; rosette leaves obovate-cuneate, 
sharply acuminate; cauline leaves more remote, alternate, the upper more 
elongated, reddish, all leaves pubescent, ciliate, entire, fleshy, 10—30 cm 
long, 8—1 3 mm broad; inflorescence corymbiform, 2—6 cm long, 4—9 cm 
in diameter, dense, many -flowered.; flowers 5-merous; pedicels ca.4— 7 mm 
long, longer than calyx; calyx pubescent, the triangular, acute sepals 
connate at base, 3—4 mm long, 1—1.5 mm broad, red, fleshy; corolla deep 
red, connate at base, the petals lanceolate, acute, pubescent on the outside, 
6—8 mm long, 1.5—2 mm broad, 2— 2. 5 times as long as calyx; stamens 10, 
red, more than half as long as corolla; follicles divergent, pubescent, 
ovoid, red, gibbous on the inside, erect, shorter than corolla; hypogynous 
scales broad, obovate, 0.3—0.5 mm long, ca. 1 mm broad; seeds oblong, 
striped, ca. 1 mm long, 0.35 mm broad. June— August. 

Dry stony slopes and rocks in the middle zone of mountains, at 
1,800— 2,000 m, to the subalpine zone.— Caucasus: W. Transc. (rarely), 
E. Transc. (Georgian SSR), mainly in S. Transc, Dag. (Derbent-Lagovskii). 
Gen. distr.: Arm. -Kurd, (former Kars Region, former Artvin District), 
Bal.-As. Min. (As. Min.). Described from the vicinity of Tbilisi. Type 
in Leningrad. 

Economic importance. Ornamental species often found in cultivation. 
Deserves attention on account of its beautiful bright red flowers. 



Section 2. EU-ROSULARIA Berger in Engl. u. Pr., Nat. Pflzfm. 18a (1930) 
465, p.p.— Perennials with fusiformly thickened underground part; 
inflorescence loose, paniculate, or corymbiform -paniculate; flowers 5-, 
6 -merous, yellow, pink, red; corolla campanulate or narrowly infundibular - 
campanulate, with cuneate base, less often calyx rounded at base. Mostly 
glandular -pubescent plants, less often glabrous. Caucasus, one species 
in Central Asia. 



Series 1 . Caucasicae A. Bor. Corolla narrowly campanulate or 
infundibular, divided to the middle or beyond. 



94 



BHHB 



3. R. sempervivum (M. B.) Berger in Engl. u. Pr., Nat. Pflzfm. 18a 
(1930) 466. — Cotyledon sempervivum M. B., Beschr. der Land. am. 
Casp. Meer (1800) 176, app., No. 46; Ej., Fl. taur. -cauc. I, 351.- Umbilicus 
sempervivum DC, Prodr. Ill (182 8) 399; Ldb., Fl. Ross. II, 175. — 
Sedum racemosum Pall, ex Ldb., Fl. Ross. II (l 844-1846) 1 75. - 

S. sempervivum Hamet in Candollea IV (1929-1931) 24 (non DC.). 

Perennial; root robust, long; entire plant minutely villous -glandular, 
5—10 (l5)cm high; rosette leaves spatulate, obtuse, truncate-rounded 
at the apex, cuneate toward base, membranously dentate-ciliate on the . 
margin; cauline leaves spatulate-oblong, obtuse, remote; inflorescence 
corymbiform -paniculate, oblong, densely glandular -pubescent, many -flowered; 
peduncles arising in axils of leaves at edge of radical rosette; flowers 
5-merous, with small lanceolate, subobtuse bracts; pedicels as long as 
calyx, calyx Y 3 — 2 / 5 as long as corolla, equal to corolla tube, pubescent, 
divided nearly to base into narrowly lanceolate, acute sepals, cuneate at 
base; corolla 8— 10 mm long, narrowly campanulate, reddish, pubescent 
on the outside, divided above the middle into 5 oblong-lanceolate acuminate, 
erect petals; stamens 10, 3 / 4 as long as corolla, with dark red reniform 
anthers; follicles lanceolate, pubescent, as long as stamens, with filiform 
beak. May— June. (Plate VII, Figure 3a). 

Stony and pebbly soils in mountains at 1,300 — 1,800 m. — Caucasus: 
S. Transc, (Mt. Alagez), E. Transc. (on Talysh border — Binotly village), 
Tal. (often). Gen. distr.: Arm. -Kurd. (Erzerum, former Kars Region), Iran. 
Described from the Caucasus. Type in Leningrad. 

4. R. radiciflora (Steud.) A. Bor. comb. nova. — U mb i 1 i c u s radici- 
florus Steud. in Kotshy pi. exs., No. 332 ex Boiss., Fl. Or. II (1872) 772.- 
U. li ba no t i c u s var. s t e u d e 1 i i Boiss., 1. c. 

Perennial; root thickened; underground part of stem short, covered by 
remnants of old leaves; flowering stems 5—15 cm high, glandular-pubescent 
in upper part, glabrous below, arising from axils of leaves on sides of 
rosette; the latter 1.5— 2.5 cm long, oblong, spatulate, more or less 
emarginate, tapering toward base, minutely glandular -hairy or subglabrous, 
membranously dentate-ciliate on the margin; cauline leaves remote, oblong, 
obtuse, glandular -pubescent; inflorescence corymbiform -paniculate, 
oblong, many-flowered, glandular -pubescent; flowers with pedicels half as 
long as calyx or else subsessile, 5-merous, with oblong -lanceolate, subobtuse 
bracts; calyx 2 / 5 — l / 3 as long as corolla, divided to base into narrowly 
lanceolate acute sepals, cuneate at base; corolla ca. 1 cm long, narrowly 
campanulate, whitish pink with darker glands, pubescent on the outside, 
divided above the middle into ovate, acute, erect petals; stamens 10, 3 / 4 as 
long as corolla, with yellow reniform anthers; follicles lanceolate, with 
filiform beak. May— August. 

Stony soils, to 3,000 m. — Caucasus : S. Transc. (Nakhichevan, Nor- 
Bayazet). Gen. distr. : Arm. -Kurd. Described from Kurdistan. Type in 
Leningrad. 

5. R.persica (Boiss.) Berger in Engl. u. Pr., Nat. Pflzfm. 18a (1930) 
466.— Umbilicus persicus Boiss., Diagn. Ser. I, 3 (1843) 14; Grossh., 
Fl. Cauc. II, 232. 



95 



Perennial; entire plant glabrous, glaucescent; flowering stems arising 
from axils of leaves at sides of rosette, 7—15 cm high; rosette leaves 
elongate-spatulate, obtuse, membranously and finely serrulate, glabrous; 
cauline leaves linear-lanceolate to linear, remote, subobtuse; inflorescence 
corymbiform-paniculate, loose, with secund, few-flowered and curved 
branches; flowers 5 -merous, campanulate, cuneate toward base; 
pedicels as long as or longer than calyx; calyx glabrous, Y 3 — l / 2 as long 
as corolla, divided to base into linear -lanceolate acute sepals; corolla 
narrowly campanulate, glabrous, pink, ca. 10 mm long, parted to V 3 into 
acute, oblong -lanceolate petals; stamens 10, adnate to l L into acute, 
191 oblong-lanceolate petals; stamens 10, adnate to l / g to corolla tube, 

shorter than corolla; hypogynous scales semiorbicular, small; follicles 
ca. 9 mm long, glabrous, elongate -lanceolate, gradually acuminate with 
a filiform beak, ca. 1 mm long; seeds oblong, brown, small, less than 1 mm 
long. June. 

Stony and dry soils, calcareous rocks, mountains, at 700—2,500 m. — 
Caucasus: S. Transc. (Ordubad, Migry). Gen. distr.: Iran., As. Min. 
Described from Iran. Type in Geneva. 

6. R. elymaitica (Boiss. et Hausskn.) Berger in Engl. u. Pr., Nat. Pflzfm. 
19a (1930) 465. — Umb i 1 ic u s elymaiticus Boiss. et Hausskn. ex Boiss. 
Fl. Or. II (1872) 770; Grossh., Fl. Cauc. II, 231 . - Sedum elymaiticum 
Hamet in Candollea IV (1929-1931) 30. 

Perennial or biennial; underground part of follicle woody, thickened, 
fusiform; flowering stems arising from axils of leaves in center of apical 
rosette; stems 7—10 (l5)cm high, glandular -pubescent, branching from 
base, forming a broad, many -flowered, thyrsoid inflorescence with secund 
branches; rosette leaves oblong-spatulate, tapering to base, obtuse, 
capitate, 1.5— 2.5 cm long; cauline leaves small, oblong-lanceolate, glandular- 
pubescent; flowers 5 -merous, as long as their drooping pedicels; calyx 
glandular-pubescent, connate at base, with oblong-ovate, subacute sepals, 
as long as or slightly more than half as long as corolla; corolla red, 
campanulate, ca. 5 mm long, connate nearly to the middle, with acuminate, 
oblong-lanceolate, erect petals, glandular -pubescent and greenish on the 
back; stamens 10, shorter than corolla, with light filaments and dark 
purple anthers; follicles lanceolate, shorter than corolla, with short beak; 
seeds small, ca. 0.5 mm long, oblong, May. 

Calcareous rocks at 2,500— 3,000 m. — Caucasus: S. Transc. (Ordubad, 
Aras River gorge). Gen. distr. : SW Iran. Described from Iran, according 
to collections of Haussknecht — Eschter, Tchinar, and Sakawa mountains. 
Type in Geneva. 

7. R. lipskyi A. Bor. sp. nova in Addenda VIII, p. 369. 

Perennial; entire plant glabrous, green, with branching, short caudex and 
thickened root; flowering stems arising from axils of leaves at sides of 
radical rosette, 5—6 cm high; rosette leaves oblong -lanceolate, 1.5—2 cm long 
4—5 mm broad, obtuse, membranous -denticulate, glabrous; cauline leaves 
oblong-lanceolate, remote, subobtuse; inflorescence paniculate, few -flowered, 
with erect branches; flowers with pedicels about as long as calyx, 5 -merous, 
.„_ campanulate, with linear bracts; calyx glabrous, V 4 — Vs a s long as corolla, 
divided to base into lanceolate, triangular, acute sepals; corolla narrowly 



96 



■^■^^^^■^■■H 



campanulate, with cuneate base, glabrous, yellow, ca. 7 mm long, parted to 
V 3 into acuminate, oblong-lanceolate petals; stamens 10, shorter than 
corolla, with yellow anthers; follicles ca. 6 mm long, glabrous, linear, 
acuminate, with filiform beak, ] / 3 as long as anthers; seeds oblong, brown, 
small, ca. 0.5 mm long. June. 

Mountains.— Caucasus: S. Transc. (Nakhichevan District — Ganza). 
Endemic. Possibly in Iran. Described from Ganza. Type in Leningrad. 



Series 2. TURKESTANICAE A. Bor. - Corolla broadly campanulate, 
with broad base, lobed to below the middle. 

8. R. hissarica A. Bor. sp. nova in Addenda VIII, p. 369. 

Perennial, with woody, long, fusiformly thickened root, with short, 
sometimes branching caudex covered with remains of old leaves, spreadingly 
villous -glandular throughout; flowering stems 5— 7 cm high; rosette leaves 
spatulate, oblong-ovate to oblong-lanceolate, cuneate at base, subobtuse, 
1—3 cm long, 3— 7 mm broad in broadest part, all leaves densely glandular- 
hairy; cauline leaves lanceolate-spatulate, obtuse, 4— 10 mm long, 1— 3 cm 
broad, remote; flowering stems arising from axils of lateral rosette 
leaves; inflorescence paniculate, loose, spreading; flowers 5 - (6)-merous, 
with pedicels longer than calyx, and with linear -lanceolate bracts; calyx 
2 / 3 as long as corolla, rounded at base, hairy, divided nearly to base into 
acute lanceolate sepals; corolla 6—7 mm long, pinkish violet, broadly 
campanulate, parted to below the middle into lanceolate, acuminate petals, 
with hairs along the midnerve on the outside; stamens 10 (12), 3 / 4 as long 
as corolla, with yellow reniform anthers; follicles lanceolate, with 
filiform apically thickened beak, longer than stamens; hypogynous scales 
linear, 1 mm long, slightly broadening toward the top; seeds 0.5 mm long, 
oblong, sulcate, brown. June. (Plate VII, Figure 2a). 

Rocks and stony soils at ca. 2,500 m. — Centr. Asia: Pam.-Al. (Gissar 
Range, Khovak; Mt. Chulbair near Sina village). Endemic. Described from 
Mt. Chulbair in the Gissar Range in Tadzhikistan. Type in Leningrad. 



Section 3. ORIENTALIA A. Bor. Perennials with tuberous, rounded 
underground parts; inflorescence long, racemose or paniculate, dense, 
many-flowered; flowers 5-, less often 6 -merous, yellow, whitish pink, 
less often red. Central Asia. 



Series 1. Racemiflorae A. Bor.— Inflorescence usually a narrow racemose 
panicle. 

9. R. paniculata (Rgl. et Schmalh.) Berger in Engl. u. Pr., Nat. Pflzfm. 18a 
(1930) 466.- Umbilicus paniculatus Rgl. et Schmalb. in A. H. P. V 
(1877) 583.- Cotyledon paniculata O. et B. Fedtsch. in Consp. Fl. 
Turk. 3 (1909) 70.- Umbelicus platyphyllus Boiss., Fl. Or. II (1872) 
773 (non Schrenk). — (?) S ed um radicosum Boiss., Diagn. Ser. 1, 10 
(1849)15.- Ic: Frod. in Acta Horti Gothoburg. VII (1932) pi. LVII; 2, textfig. 
90 — 98 (sub S. radicosum Boiss.). 



97 



Perennial; root tuberous, rounded -oval, with a bundle of slender roots; 
stems 20—35 (40)cm high, erect, simple, glabrous, leafy, arising from axils 
of rosette leaves; rosette leaves flat, spatulate, obovate or oblong, mostly 
acuminate, fleshy; cauline leaves remote, alternate, sessile, spatulate, oblong 
or lanceolate, acute, the lower 1— 1.5 cm long, diminishing upward; 
inflorescence a racemose, elongated panicle, ca. 10—15 cm long, 2—3 (4)cm 
in diameter, with numerous short, recurved, divaricate branches; 
flowers 1—3 (sometimes more) on each branch, 5 -merous, with small, 
ovate-lanceolate, acute bracts; pedicels from shorter to longer than flowers; 
calyx divided into ovate -lanceolate, acute sepals, ca. x / 3 as long as corolla; 
petals ca. 5—6 mm long, connate from base nearly to the middle or lower, 
lanceolate, acuminate, divergent, pale yellow (dry), greenish or pinkish, 
with dark nerves; stamens 10, shorter than petals, with pale yellow filaments 
and rounded, darkish anthers; hypogynous scales obtuse, subquadrate; 
follicles erect, ovoid -lanceolate, the filiform beak 2—3 cm long, longer than 
petals; seeds small, less than 1 mm long, brown, ovoid. June — July. 

Stony slopes at 2,000-3,000 m. - Centr.Asia: Mtn. Turkm. (Kopet Dagh 
Range), Pam.-AL, T. Sh. (W. part). Gen. distr. : Iran. Described from 
Zeravshan. Type in Leningrad. 

Note. Highly polymorphic species; owing to lack of material, it 
is as yet impossible to subdivide the species. Transcaspian specimens 
differ somewhat from the Pamir -Alai plants and may belong to an independent 
species. (S e d u m radicosum Boiss.). 

10. R. subspicata (Freyn et Sint.) A. Bor. comb. nova. — U. subspicatus 
Freyn et Sint. in Bull. Herb. Boiss. (1906) 195.— Cotyledon subspicata 
Fedtsch. O. et B. in Consp. Fl. Turk. 3 (1909) 69. 

Perennial, tuberous root; green, glandular -pubescent; flowering stems 
arising in radical rosettes, curved at base; rosettes ca. 6 cm in diameter; 
radical leaves sessile, flat, oblong, 23—30 mm long, 5—6 mm broad, acute, 
entire, pubescent on the margin, otherwise glabrous; cauline leaves 
1.7—7.5 mm long, erect, terete, sessile, lanceolate -oblong, acute, sparsely 
glandular; flowers in long raceme, with acute lanceolate -subulate bracts, 
7 mm long, 0.5 mm broad; pedicels 1 — 1.5 mm long, erect, elongated, about 
half as long as calyx, glandular -tuberculate; calyx with oblong-elliptic, 
glandular sepals, 3 mm long, 1 mm broad, half as long as corolla; corolla 
campanulate, yellow, connate to % of its length, the petals glabrous, elliptic- 
oblong, acute or short acuminate, red along the midnerve, 6 mm long, 
1.5 mm broad; stamens shorter than petals, with elliptic, dark anthers, 
yellow pollen, and white filaments, pistils lanceolate, with filiform, erect 
styles, slightly longer than petals. June. 

Among Juniperus in mountains. — Centr. Asia: Mtn. Turkm. (Kopet 
Dagh Range, near Solyukli village), Pam. -Al. (near the Zirabulak Railroad, 
Shakhrisyabs). Gen. distr.: possibly in adjacent parts of Iran. Described 
from the Kopet Dagh Range, from the vicinity of Solyukli. Type in 
Leningrad. 

Note. Specimens from Pamir -Alai are distinguished from the Kopet 
Dagh R. subspicata by their greater pubescence and by their branching, 
many -flowered inflorescence. 

Series 2. Thyrsiflorae A. Bor. — Inflorescence broad, paniculate. 



98 



nowmBHH^HH 



125 




PLATEVII 1-Penthorum chinense Pursh: a) flower, b) fruit; 2 - Rosula r U h 1 »»''"? 
PLATE VU. i reniuuiu. crhischkinii A. Bor.: a) tlowcr, 

a) flower; 3 - R.semperv ivum fM.B.) Berger: a) flower; 4 - R. sch is 

b) rosette leaves; 5 - R. p la t y ph y 11 a (Schrenk) Berger: a) flower. 



99 



11. R. glabra (Rgl. et Winkl.) Berger in Engl. u. Pr., Nat. Pflzfm. 18a 
(1930) 466.- Umbilicus glaber Rgl. et Winkl. in A. H. P. VI (1879) 
302.- Cotyledon glabra Fedtsch. O. et B. in Consp. Fl. Turk. 3 (1909) 
70.- Cotyledon ferganica Drob. in Sched. — Ic. : Gartenfl. (1880), 
tab. 1019, f. 1. 

Perennial; roots arising in a bundle from rounded underground part; 
entire plant glabrous; stems 1 — 3, suberect, arising at sides of radical 
leaf rosette, 6—25 cm high, leafy; leaves fleshy, flat; rosette leaves 
spatulate -elliptic, the outer subobtuse, the inner short -acuminate, all 
leaves deep green, elongated to (3)4—6 cm, sparsely and papillose- 
hispidulous; cauline leaves remote, alternate, the lower elliptic -oblong, the 
upper oblong, gradually passing into linear bracts; inflorescence compound, 
paniculate, many -flowered, spreading, 4— 7 cm in diameter, long scorpioid, 
with secund branches; flowers remote, short pediceled, 5-merous; calyx 
divided nearly to base into ovate, acute sepals, ! / 4 — Y 3 as long as corolla; 
corolla campanulate, yellow or yellow -green, parted nearly to the middle 
into elliptic, acute, erect, subcarinate petals 5— 8 mm long; stamens about 
as long as corolla, with yellow anthers and filaments; follicles glabrous, 
ovoid, longer than petals, with beak ca. 2 mm long. May —June. 

Dry stony slopes at 1,500— 3,000 m. — Centr.Asia: Pam.-Al. (Pamir, 
Darvaz, Alai Range, Shurabad [?] district). Endemic. Described from 
E. Turkestan (E. Regel). Type lost ? 

Note. According to Regel, the length of the flower is 8 mm, but in fact 
it is 5— 8 mm; in the Gartenflora drawing, the stamens protrude too far. 
Apparently the Gartenflora illustration was drawn after a R. glabra type 
which has not been preserved. 

12. R. turkestanica (Rgl. et Winkl.) Berger in Engl. u. Pr., Nat. Pflzm. 
18a (1930) 466.- Umbilicus t u r k e s t a ni c u s Rgl. et Winkl. in A. H. P. 
VI, 2 (1879) 301.- Cotyledon turkestanica Fedtsch. O. et B., Consp. 
Fl. Turk. 3 (1909) 69.- Cotyledon dubius C. Winkl. in sched.- Ic. : 
Gartenflora (1880) tab. 998, f. 1. 

Perennial; underground part tuberous; stems 7— 10 cm high, arising 
at side of radical leaf rosette, glabrous, greenish or with reddish dots, 
few, ascending, leafy; radical leaves flat, in a dense rosette, the outer 
lanceolate or oblong-lanceolate, 2—3 cm long, acuminate, the inner 
spatulate -ovate, acute, all leaves hispidulous-ciliate on the margin, with 
both surfaces hispidulous-ciliate and later glabrous; inflorescence 
paniculate, 5—15 cm long, with subscorpioid secund branches, many- 
flowered, densely flowered; flowers 5 - or 6 -merous, with pedicels as 
long as or longer than calyx; calyx divided nearly to base into lanceolate 
or elliptic -lanceolate, acute sepals, % — Y 3 as long as corolla; corolla 
campanulate, purple, drying yellowish with violet tips, parted to the 
middle into declinate, lanceolate, acute petals, ca. 4 mm long; stamens 
10— 12, about as long as corolla; follicles oblong-lanceolate, erect, united 
nearly to the apex, pubescent at base, with short, filiform beak. June. 

Gravelly slopes. — Centr. Asia: T. Sh. Endemic. Described from 
E. Turkestan. Type unknown. 

Note. Polymorphic species requiring detailed study, which is impossible 
owing to lack of material. The type may be surmised from the Gartenflora 
drawing. 



100 



H 



Section 4. CAMPANELLA A. Bor. - Genus Sedum Group 
Umbilicoides Frod. in Acta Horti Gothoburg. VI (1931 ) 25 p. p. — 
Perennials with tuberous underground parts; inflorescence short, 
corymbiform, corymbiform-paniculate or umbellate, loose; flowers 
5—8 -merous, white, yellow -violet, pinkish or violet, less often yellow; 
corolla broadly campanulate, with broad base. Plants glabrous or 
puberulent or ciliate only on leaf margin. Central Asia. 



Series 1. Luteae A. Bor.— Flowers yellow. Small plants, 3—5 cm high. 

13. R. kokanica (Rgl. et Schmalh. ) A. Bor. comb. nova. — Sedum 
kokanicum Rgl. et Schmalh. ex Rgl., Descrip. plant, nov. in Izv. o-va lyub. 
estestv. XXXIV, 2 (1882) 26; A. P. Fedchenko, Puteshestvie v Turk in 

1 8 (l 881 ), No. 64. - Ic. : Fr5d. in Acta Horti Gothoburg. VI (l 931 ) pi. XV. 

Perennial or biennial; roots thickened, fleshy, short; entire plant ca. 
3 cm high; stems small, erect or ascending, mostly numerous, arising 
from radical leaf rosette, glabrous, with sparse bristles near tip, densely 
leafy; leaves fleshy, flat, hispid -ciliate only on the margin, glabrous 
on the surface; radical leaves in 9 dense rosettes, lanceolate, acuminate; 
cauline leaves ovate -lanceolate to lanceolate, acuminate; inflorescence 
few-flowered, corymbiform, subcapitate; flowers 6 -merous, 3 mm long, 
with very short pedicels and with linear bracts; calyx half as long as 
corolla, with ovate -lanceolate, acuminate, triangular, 3 -nerved sepals; 
petals narrowly lanceolate, short~connate at base, acute, 1 -nerved, yellow, 
3 mm long; stamens 12, those opposite to petals V 3 as long as them, with short 
filaments, adnate to petals to half their length, the stamens opposite sepals longer 
than the other 6, half as long as petals; anthers large, oblong-ovate, 
yellow. June. 

High mountains, at 2,800—3,500 m. Known and described from a single 
site.— Centr.Asia: Pam. -Al. (near Kokand, Shchyrovskii Glacier). Type 
in Leningrad. 

14. R. lutea A. Bor. sp. nova in Addenda VIII, p. 370. 

Perennial; root tuberous, rounded -oval, 0.6—2 cm thick, giving rise to 
several thinner roots; stems 3—5 cm high, 2—3, arising in leaf rosette, 
glabrous, with remote leaves;' leaves fleshy, flat, the radical in a dense 
rosette, oblong-lanceolate, subobtuse, glabrous, with very sparsely ciliate 
margin or eciliate; cauline leaves oblong-lanceolate, subobtuse; 
inflorescence many -flowered, corymbiform-umbellate, rather loose; 
flowers 5- or 6 -merous, with long, slender pedicels; sepals 1/3 as long 
as corolla, ovate, acute, ca. 2 mm long; petals oblong-lanceolate, short- 
connate at base, mucronate, 1 -nerved, yellow, 5—7 mm long; stamens 10—12, 
those opposite to sepals slightly shorter than and those opposite to petals 2 / 3 
as long as petals, with oblong -ovate dark violet anthers and yellow 
filaments; hypogynous scales semiorbicular, truncate, very small; 
follicles lanceolate, many -seeded, with filiform beak ca. 1 mm long. July. 

Calcareous rocks, thickets of woody -shrubby vegetation.— Centr.Asia: 
Pam.-Al. Endemic. Described from E. Tadzhikistan (Vakhsh Range, near 
Kamoli village). Type in Leningrad. 



101 



Series 2. Alpestres A.Bor.— Flowers white, violet, or yellow -violet. 
Plants (5)8-15 (18) cm high. 

15. R. alpestris (Kar. et Kir. ) A. Bor. comb. nova. — Umbilicus 

alp est r is Kar. et Kir. in Bull. Soc. Nat. Mosc.XV (1842) 354.- Cotyledon 
alpestris Fedtsch. O. et B., Consp. Fl. Turk. 3 (190 9) 70.- Sedum 
umbilicoides Rgl. in A.H. P. V, 1 (l 877) 263; Gartenflora (1877) 2 90; 
Fedtsch. O. et B., Consp. Fl. Turk. 3 (l 909) 73.- S.olgae Rgl. et Schmalh. 
ex Rgl., Descrip. plant, nov. in Izv. o-va lyub. estestv. XXIV, 2 (1882) 26. — 
S. acuminatum Hamet in Candollea IV (192 9-1931) 2 3.- Ic: Gartenflora 
(1877) tab. 917; Frod. in Acta Horti Gothoburg. VI (1931 ) pi. XVI. 

Perennial; root rounded -tuberous; stems arising from axils of rosette 
leaves, erect or ascending, leafy, glabrous, 5—8 (l2)cm high; leaves fleshy, 
flat, hispidulous-ciliate on the margin only at the apex, otherwise glabrous; 
basal leaves rosulate, 1.5—2.5 cm long, 3—6 mm broad, oblong-lanceolate 
or oblong, acuminate; rosettes 1.5 — 3 cm in diameter; cauline leaves 
sessile, oblong or oblong-lanceolate, acuminate; inflorescence simple, 
few-flowered, corymbiform or corymbiform -paniculate or racemose 
flowers 6— 8-merous, with pedicels shorter than corolla, or upper flowers 
with elongated pedicels, with small ovate -lanceolate bracts; sepals 
lanceolate, acute or acuminate, glabrous, 3 -nerved, half as long as corolla; 
petals connate at base, lanceolate, acute, 3 -nerved, white or pinkish, along 
the keel violet or reddish, recurved and divergent at the apex, 6—9 mm long; 
stamens 12—16, shorter than petals, with ovate, dark anthers and light 
filaments; hypogynous scales small, semiorbicular, truncate or rounded 
at the tip, entire; follicles many -seeded, convergent, with filiform beak 
ca. 1 mm long, glabrous; seeds small, brown, ovoid, less than 1 mm long. 
June — July. 

Stony soils at 1,500— 3,000 m. — Centr.Asia: Dzu. -Tarb. (Dzungarian 
Ala-Tau), T.Sh., Pam.-Al. (Alai Range). Gen. distr. : Dzu.-Kash. 
(Kuldja). Described from the Dzungarian Ala-Tau. Type in Leningrad. 

Note. In his description of S. o lg a e, Regel cites only 6 -merous 
flowers, but in fact flowers with 7 petals occur in authentic specimens; 
this is true also of S. umbilicoides Rgl., for which 7 -merous flowers 
are recorded, though 6 -merous flowers are not exceptional. Umbilicus 
alpestris Kar. et Kir. also has 6 - and 7 -merous flowers, but they are 
more united at the base than in S. o 1 g a e and S. umbilicoides. The 
number of stamens (8— 10 ) indicated by Regel for S.olgae is erroneous; 
this species always has a double number of stamens, viz., 12 — 14. Thus, these 
three species cannot be differentiated on the basis of the material available. 
The specimens collected in the Ketmen Range are distinguished by their 
stockiness, few flowers (often 1 ), and more intense color of flowers. Certain 
specimens from the Kirghiz Ala-Tau (collections of Massagetov and 
Massal'skii) and from the Beishmunak natural boundary area (collections 
of Taranenko) are distinguished by linear-lanceolate leaves. 

16. R. schischkinii A. Bor. sp. nova in Addenda VIII, p. 370. 
Perennial; roots tuberous; stems 1—7, arising in axils of lateral 

leaves of radical rosette, glabrous, 12—20 cm high, leafy, erect or ascending; 
leaves fleshy, flat; rosette leaves hispidulous, densely white-ciliate- 
margined, especially at the apex, obovate -oblong, acute, 1.5—2.5 cm long, 
0.5—1 cm broad; rosette 3— 8 cm in diameter; cauline leaves oblong, hispid 



102 



HH 



on margin and blade surface, acuminate, remote, ca. 0.7—1 .5 cm 
long, ca. 0.2—0.4 cm broad; inflorescence corymbiform -paniculate, many- 
flowered; flowers 5— 7-merous, long-pediceled, with small, linear, acute 
bracts; calyx with ovate, 3 -nerved, subacute sepals, 2.5—3 times as long 
as corolla, with long white, sometimes deciduous, hairs at base; petals 
subobtuse, apiculate, oblong-lanceolate, 3 -nerved, white or pinkish, drying 
dark violet along the keel, connate at base, initially divergent, later erect, 
0.8—1 cm long; stamens 12 (14), shorter than petals; filaments white, 
dilating toward base; anthers ovate; hypogynous scales small, semiorbicular; 
follicles pubescent dorsally, convex ventrally, with subulate, reflexed beak more 
than 1 mm long, the follicles not longer than petals; seeds ca. 1 mm long, 
oblong. July- August. (Plate VII, Figure 4— 4b). 

Grassy slopes in the steppe zone at ca. 2,000 m. — Centr.Asia: T. Sh. 
(E.part — Sarydzhaz, Kumurchi, Lake Issyk-Kul). Endemic. Described 
fromSarydzhaz. Type in Leningrad. 

Note. Closely related to R. a lp e st r i s (Kar. et Kir.) A. Bor., from 
which it is distinguished by its larger size, shape of rosette leaves, 
pubescence of leaves, calyx and fruit, mostly 5- (rarely 6 - or 7-merous) 
flowers. 

17. R. tadzhikistana A. Bor. sp. nova in Addenda VIII, p. 371. 
Perennial; roots tuberous only in upper part, cordlike below, few; stems 

3—6 (8) arising in axils of rosette leaves, erect or ascending, leafy, glabrous, 
5 — 10 (15) cm long; leaves fleshy, flat, glabrous; radical leaves rosulate, 
the young ones white-setose, long ciliate throughout margin, later 
glabrescent, 0.5— 0.8 cm broad, 1—1 .5 cm long, ovate to oblong, mucronate; 
leaf rosettes (2.5) 3— 5 cm in diameter; cauline leaves sessile, oblong- 
lanceolate, acute, entire, remote; inflorescence simple, umbellate - 
corymbiform, composed of (2)5—10, sometimes more flowers, usually 
3-branched; flowers 6 - or 7-merous, as long as or longer than their 
pedicels, with linear bracts; sepals ovate, acute, glabrous, / 4 — l / 3 as long 
as corolla; petals connate at base, lanceolate, subobtuse or subacute, 
3 -nerved, yellow -red in the middle mostly dark lilac, with yellow edges, 
5—7 (8) mm long; stamens 12 — 14, 2 / 3 as long as petals, with yellow filaments 
and rounded yellow, later dark anthers; hypogynous scales small, semi- 
orbicular, entire; follicles many-seeded, lanceolate, with erect subulate 
beak, ca. 1 mm long; seeds small, ca. 1 mm long, yellow, oblong -ovoid, 
subacute. July— August. 

Stony slopes, rock crevices, taluses in the high-mountain zone, and 
alpine meadows. — Centr. Asia : Pam. -Al. (Shugnan, Pamir, Darvaz, Roshan, 
Obi-Khingou River basin). Endemic. Described from Dasht-i-Uzbekon. 
in E. Tadzhikistan. Type in Leningrad. 

18. R. platyphylla (Schrenk) Berger in Engl. u. Pr., Nat. Pflzfm. 18a (1930) 
466. — U mb i 1 i c u s platyphyllus Schrenk in Fisch. et Mey., Enum. 

pi. nov. (1841) 71; Ldb., Fl. Ross. II, 175; Boiss., Fl. Or. II (1872) 773, p.p. -? 
U.pulvinatus Ost.-Sack. et Rupr., Ser. tiansch. (1869) 47.— Cotyledon 
platyphylla O. et B. Fedtsch., Consp. Fl. turk. Ill (1909) 69.- Ic: 
Gartenflora (1880) tab. 998, f. 2. 



103 



Perennial; underground part of root thickened, tuberous, rounded -oval, 
with few thick roots from which arise small roots; flowering stems 
few (1—4), 10—15 cm high, ascending, simple, puberulent, arising at edge 
of radical leaf rosette; rosettes 5 — 10 cm in diameter; rosette leaves 
flat, rhomboid -ovate or spatulate, ca. 1.2—2 cm broad, 1.5—2.5 (4) cm long, 
obtuse or concave or bluntly acuminate, sometimes tapering toward base, 
ciliate -margined, puberulent on both surfaces; cauline leaves remote, 
alternate, sessile, oblong, 1 — 1 .5 cm long, 0.4— 0.5 cm broad, ciliate-margined, 
puberulent; inflorescence corymbiform, short, ca. 3—4 cm in diameter, 
3—5 cm long, with glandular -puberulent scabrous branches, many -flowered, 
the pedicels shortex- than flowers, the branches long, secund, with small, 
oblong-linear bracts; flowers 5-merous; calyx with obtuse, ovate sepals, 
Y 3 — '/ 2 as long as petals; corolla white, drying pale yellow, 5—7 mm long, 
with tube as long as calyx; limb of corolla 2 — 3 times as long as tube, 
with 5 subobtuse, apiculate, ovate petals; stamens 10, shorter than corolla, 
with yellow anthers and white filaments; follicles ovoid -oblong, with 
filiform beak ca. 2 mm long; seeds brown, less than 1 mm long, oblong - 
ovoid. June— July. (Plate VII, Figure 5a). 

Stony and gravelly slopes, rock crevices, occurs singly. — Centr. Asia: 
T. Sh., Dzu.-Tarb. Gen.distr.: Dzu. -Kash., Kuldja. Described from the 
Dzungarian Ala-Tau. Type in Leningrad. 

Note. The stamens are incorrectly represented in the Gartenflora 
drawing. 



Subfamily 5. PENTH0R0ID2AE (ENGL.) A. Bor. - Family Saxifragaceae, 
subfamily Penthoroideae, tribe Penthoreae Engl. u. Pr. Nat. Pflzfm. 18a 
(1930) 112.- Family Penthoraceae Van Tieghem in Journ. bot. XII (1898) 
150; Rydberg in Britton North American Fl. XXII, 1 (1905) 75.- Follicles 
united nearly to the middle, dehiscing by a ring of transverse clefts; 
flowers 5-merous; petals inconspicuous or absent. Nonsucculent plants. 



Genus 703. PENTHORUM GRONOV. 

Gronov.ex L., Coroll. gen. (1737) 8; L., Sp.pl. ed. 1 (1753) 432. 

Flowers 5- or 6-, rarely 7 - or 8-merous, small, greenish; calyx 
5 -merous, green, persistent; petals 5 - or 6, green, inconspicuous or absent; 
stamens 10 — 12, with filiform filaments and oblong anthers, dehiscing by 
longitudinal clefts; ovaries 5 or 6, connate at base, or nearly to the middle, 
with short styles and capitate stigma; follicles unilocular, compressed, 
arranged on a pyramidal receptacle and connate to half their length; seeds 
numerous, small, ovoid or oblong, minutely tuberculate -glandular. 
Perennial plants; leaves not fleshy, alternate, lanceolate or oblong- 
lanceolate, unequally serrate; inflorescence terminal, corymbiform, 
composed of (l ) 3 — 1 secund branches, scorpioid at the tip. 

1. Plants 40—60 (85) cm high; stems solitary, simple, less often branching; 
leaves elongate-linear-lanceolate, 5—10 cm long, 1 — 1.5 cm broad; 
inflorescence corymbiform, many -branched; flowers 3— 5 mm long 
1 . P. chinense Pursh. 



104 



BHZHmn 



133 



+ Plants 15— 20 cm high; stems numerous or solitary, mostly branching; 
leaves oblong-lanceolate or lanceolate, 1—4 cm long, 0.4—0.7 cm 
broad; inflorescence composed of solitary branches; flowers 2—3 cm 
long 2. P. humile Rgl. et Maack. 

1. P. chinense in Flor. bor. amer. I (1814) 323 inobs.; DC, Prodr. Ill, 
414; Kom., Fl. Manch. II, 403; Kom. and Alis., Opred. rast. Dal'nevost. kraya I, 
602.- P. intermedium Turcz. in Bull. Soc. Nat. Mosc. VII (1837) 152, 
No. 82.- Ic: E. Regel, 1. c.,t. VI, f. 1-4. 

Perennial; rhizome short, vertical; stems solitary, erect, simple, less 
often branching at base, 40—60 (85) cm high, densely leafy, glabrous at base, 
glandular -puberulent at summit; leaves remote, elongate -linear -lanceolate, 
usually glandular on the margin, unequally and acutely serrulate, acute, not 
fleshy, glabrous, green on both sides, erect, 1 — 1.5 cm broad, 5—10 cm long, 
cuneate at base, short -petioled; inflorescence corymbiform, terminal, 
composed of 3—10 scorpioid branches, few -flowered or many -flowered, 
with short sparse glands on branches; flowers subsessile, with short 
glandular -pubescent pedicels, 5-, rarely 6— 8-merous; calyx broadly 
campanulate, connate at base, with 5 or 6 ovate -lanceolate, acute sepals; 
petals almost always absent; stamens 10—12 (14—16), arranged in 2 series, 
5—6 (8) stamens shorter than or as long as sepals and 5 longer than sepals; 
anthers oblong; follicles 5 or 6, rarely 7 or 8, connate to the middle, with 
stellately spreading, thickish beak; seeds ovoid -oblong, very small, sharply 
tuberculate by small glands. Fl. July — August, fr. August — September. 
(Plate VII, Figure la-b). 

Along rivers, on silty bars, occasionally along roadside ditches; has 
recently become a component of ricefield weeds.— Far East: Uss. 
Gen. distr. : Jap.-Ch. (China, Korea, Japan). Described from China. 

2. P. humile Rgl. et Maack ex E. Regel in Mem. Ac. Sc. Petersb. VII, 
ser., IV, No. 4 (1861) 64.- Ic: Regel, 1. c.,t. VI, f. 5-8. 

Perennial; rhizome long, branching or nearly branching, creeping or 
obliquely ascending; stems numerous, simple or branching, 15—20 cm high, 
densely leafy, sulcate, glabrous or with small glandular hairs at the 
summit; leaves alternate, oblong-lanceolate or lanceolate, unequally 
serrulate and glandular on the margin, not fleshy, green, ca. 0.4—0.6 cm 
broad, 1—4 cm long, tapering to a short petiole or subsessile, acute, 
glabrous; inflorescence terminal, few-flowered, with solitary, glandular- 
pubescent scorpioid branches; flowers small, 2 — 3 mm long, 5 -merous, 
subsessile, with very short, glandular -pubescent pedicels; calyx connate 
at base, with 5 ovate -lanceolate, acute teeth; corolla absent; stamens 10 
slightly longer than calyx; follicles 5, with short, thick beaks, stellately 
spreading at maturity, connate to the middle. August. 

Moistened and silty soils, near water, in river valleys.— Far East: 
Uss. (Suifun River near the city of Voroshilov [Ussuriisk]). Endemic. 
Described from S. Manchuria, from the Sungacha River. Type in Leningrad. 

Note. Species very closely related to P. chinense Pursh; this may be 
a suppressed form of the described species with the same habitats. 
134 R. sedoides L., distinguished by lanceolate leaves more than 3 cm 
broad, grows in moist forest plots in the Atlantic Coast states of North 
America (Florida, Louisiana and Texas, from Ontario to Wisconsin). 



105 



Family LXXIV. SAXIFRAGACEAE DC. 

Flowers small or of medium size, cyclic, usually differentiated into 
calyx and corolla, sometimes, owing to reduction of petals, with uniseriate 
perianth, mostly 5-merous, less often or up to 10 -merous, bisexual, less 
often unisexual, mostly actinomorphic, occasionally with a tendency toward 
zygomorphy. Receptacle convex, flat, or, if concave, then ovaries adnate 
to receptacle in lower part to throughout their length. Stamens mostly 
obdiplostemonous or haplostemonous, less often numerous. Carpels 
usually connate, fewer than petals, less often carpels free and as many 
as petals. Styles usually free. Ovary mostly bilocular, sometimes 
unilocular, occasionally 5 -locular; ovules varying in number, multiseriate, 
parietal, the placenta often inflated. Seeds small, numerous, with abundant 
endosperm and small erect embryo. — Herbs or woody plants, with usually 
alternate, rarely opposite leaves; usually exstipulate. 

1 . Herbaceous plants 2. 

+ Woody plants 7. 

2. Flowers 5 -merous, differentiated into calyx and corolla 3. 

+ Flowers 4 -merous, with calyx only 708. Chrysosplenium L. 

3. Petals pinnatifid, with very narrow lobes 707. Mitella L. 

+ Petals entire 4. 

4. Perennials, large, with 2— 3 -fid leaves; flowers small, in a dense, 
many-flowered panicle 704. Astilbe Hamilt. 

+ Perennials, less often annuals, with entire, spatulate or palmatifid 

leaves; flowers solitary or in variously shaped few- or many -flowered 
inflorescences 5. 

5. Flowers solitary; 5 stamens alternating with 5 glandular-ciliate 
staminodes 70 9. Parnassia L. 

+ Flowers usually in an inflorescence, less often solitary, with 10 

stamens, without staminodes 6. 

6. Leaves large, to 30 cm long, covered with sunken multicellular 

glands (punctate when magnified); flowers lilac -red 

705. Bergenia Moench. 

Leaves not more than 10 cm long, without sunken glands; flowers white, 
greenish, yellow, or reddish 706. Saxifraga L. 

7. Leaves opposite 8. 

+ Leaves alternate 9. 

8. Shoots usually unarmed; flowers in normal racemes; pedicels 
sometimes very short; ovary short -stalked, often inconspicuously .... 
713. Ribes L. 

+ Shoots covered with prickles; flowers in 1—3 (5 )-flowered, fascicular 
racemes; pedicels not developing; ovaries with long stalk persistent in 
fruit 714. Grossularia Mill. 

9. Plants with stellate hairs on shoots, leaves, and receptacle 

711. Deutzia Thumb. 

+ Plants without stellate pubescence 10. 

10. Flowers in simple, few -flowered racemes, bisexual, large 

710. Philadelphus L. 

+ Flowers in pyramidal or corymbiform many -flowered panicles, the 

marginal sterile, large, the others bisexual, small . . . . 712. Hydrangea L. 



106 



^■■B 



Subfamily 1. SAXIFRAGOIDEAE A. Bor. in Aschers Fl. Prov. Brandenb. I 
(1864)61.— Flowers usually 5-, less often 4 -merous, differentiated into 
calyx and corolla or with calyx only. Carpels 2, rarely 4, free or connate 
in lower part. Ovules with 2 integuments. Herbs with alternate leaves; 
leaves exstipulate or with stipular sheath excrescences. 

Tribe I. SAXIFRAGEAE DC. Prodr. IV (1830) 17.- Carpels usually 
connate, with free style, less often carpels free. 



Genus 704. ASTILBE HAMILTON 

Hamilton in Don, Prodr. Flor. Nepal. (1825) 910.— Hoteia Morr.et Dene in Ann. Sc. Nat. Paris Ser. 2, 
II (1834) 316, tab. II. 

Calyx campanulate, with 5, less often 4 free sepals; petals 4, 5, or none; 
stamens 8—10, with long filaments and geminate, cordate anthers; ovary 
2 — 3-locular, with many ovules, divided to base into 2 or 3 lobes with 2 
subsessile low stigmas; capsule bilobate (rarely trilobate). Large 
herbaceous perennials with simple stems and delicate 2 — 3 -fid dentate 
leaves; flowers numerous, in terminal racemose inflorescences. 

1. A.chinensis (Maxim.) Franch. et Sav., Enum. PI. Japon. I (1875) 
34; Kom., Fl. Manch. II (l 904) 407.- A.odontophylla Mlq. Ann. Muss. 
Bot. Lugd. Bot. IF (186 7) 96. - Hot eia chinensis Maxim., Prim. Fl. 
amur. (1859) 120.— H.thunbergii Regel, Tentam. Fl. Ussur. in Mem. Acad. 
Sc. VII, ser., IV (1861 ) 68 (non Sieb. et Zucc). - Ic: Gartenfl. XII (1863) 
tab. 389; LV (l 916) tab. 1546 ; Bot. Mag. tab. 7880. 

Perennial; rhizome woody, dark brown; stem to 1 m high, simple, 
solitary, erect, lignifying in lower part, smooth, glabrous or sparsely 
pubescent by dark rufous hairs, leafy; radical leaves 3 or 4, the blade 
shorter than the petiole, glabrous or pubescent, sulcate in upper part; 
petioles of cauline leaves short, with articulate, rufous, ovate stipules 
at base; all leaves lustrous above, dull below, covered on both sides with 
small rufous bristles, mainly along veins and margin, compound, bipinnate, 
the leaflets oval or oval-lanceolate, with cuneate or slightly cordate base, 
short -acuminate, doubly subulate -dentate on margin, sessile, only the 
terminal leaflet petiolate; inflorescence 10— 30 cm long, terminal, racemose, 
with erect lateral branches; peduncles puberulent; pedicels very short, 
hairy, with small membranous bracts; calyx with broadly ovate, acute, 
entire, white scarious sepals 1 — 1.5 mm long; petals 5, violet, 3—5 times as 
long as calyx, narrowly linear, 1—3 -nerved, subfiliform toward base, short - 
acuminate; stamens 10, longer than calyx, with violet filaments; ovaries 
pale brown, the upper ovary bilocular, sharply divided to base into 2 lobes, 
with 2 small subsessile stigmas; capsule light brown, with divergent lobes, 
dehiscing by ventral suture, with 15—20 curved, narrowly elliptic, acuminate 
seeds 0.5—1 mm long. 

Open broadleaf forests, forest edges, among shrubs.— Far East: Ze.-Bu. , 
Uss. Gen. distr. : Jap.-Ch., Mong. Described from the lower reaches of the 
Amur. Type in Leningrad. 

* Treatment by A. S.Lozina-Lozinskaya. 



107 



Genus 705. BERGENIA MONCH. 

Mo'nch in Method. Plant. (1794) 664. 

Flowers 5-merous; calyx fused at base, not adnate to ovary; corolla 
polypetalous, campanulate; petals red, pink, or white, unguiculate; stamens 
10; ovary 2- or 3-locular, with 2 or 3 styles. Stems leafless; inflorescence 
crowded, corymbiform; leaves rosulate, petiolate, large, coriaceous.* 

1. Petals cuneate, tapering to long claw; leaves ovate, cuneate or 

subtruncate at base 2. B.pacifica Kom. 

+ Petals with a short, broad claw; leaves rounded -elliptic or broad - 

ovate, rounded or cordate at base 1. B. crassifolia (L. ) Fritsch. 

1. B. crassifolia (L.) Fritsch. in Verh. Zool. Bot. Ges. XXXIX (1889)575; 
Kryl., Fl. Zap. Sib. VI, 1430.- B.bifolia Moench, Meth. PL (1794) 664.- 
Saxifraga crassifolia L., Sp. pi. (1753) 401 ; Ldb., Fl. Ross. II, 214. - 
Ic. : Bot. Mag. tab. 1 96. — Vernacular names: salai, kachintsy, koyashan 
(Turkic). 

Perennial; rhizome thick, creeping; stem thick, leafless, 6—50 cm high, 
glabrous; radical leaves in dense rosette, petiolate, lustrous, glabrous, 
puncticulate below, wintering, the blade broadly elliptic or rounded, less 
often broadly obovate, rounded at base or cordate, obscurely dentate, 3—35 cm 
long, 2.5—30 cm broad, the broad petioles not longer than the blade, with 
membranous sheathing stipules at base; flowers ebracteate, in a terminal, 
dense, paniculate -corymbiform inflorescence, usually in pairs with long 
reddish pedicels to 4 cm long, elongating in fruit to 10 cm; calyx 
campanulate, dissected to the middle into oval, apically rounded sepals to 
4 mm long, glabrous; petals obovate or broadly ovate, with broad, short 
claw 10—12 mm long, 6— 8 mm broad, with obtuse -rounded apex and many 
nerves, lilac -red; stamens 10, twice as long as calyx and shorter than 
style; ovary half-inferior; capsule with 2 divergent lobes, dehiscing 
along ventral suture; seeds numerous, smooth, glabrous, 1.5—2 mm long. 
Fl. May - July, fr. August. 

Rocks, stony slopes, rock streams, old moraines, in the forest and alpine 
zones. W.Siberia: Alt.; E.Siberia: Ang. -Say. , Yenis., Dau., Lena -Kol.; 
Centr.Asia: Dzu. -Tarb. Gen. distr»: N. Mong. Described from Siberia. 
Type in London. 

Economic importance. Rhizome and leaves contain a great quantity 
of tannin; the plant is used for tanning. 

2. B. pacifica Kom. in Fedde, Repert. sp. nov. IX (l 91 1 ) 393. — B. crassi- 
folia var. pacifica Nekrassova in Fl. Az. Ros. in 11 (1917) 15. — Ic. : 
Kom. and Alis., Opred. rast. Dal'nevost. kraya I (l 931 ) Plate 181. 

Perennial; rhizome horizontal, long, thick, 30—50 cm long, not branching; 
leaves glabrous, thick, firm, light green, lustrous, with prominent midrib, 
elliptic, slightly cuneately tapering downward, or subtruncate at base, 
rounded at the apex, slightly dentate-sinuate, 4—15 cm long, 3—9 cm broad; 
petioles shorter than the blade, glabrous, with amplexicaul, membranous, 
broad stipules; flowering stem scapiform, glabrous, reddish, ribbed, to 

* B.gorbunovii B. Fedtsch. (Trudy Tadzh. bazy AN II, 1936, 141),, a little-known species, is distinguished 
from those cited here by its coarsely ciliate- margined leaves; flowers and fruit unknown.— Centr.Asia: 
Pam.-Al. (known only from Yazgulem). Type in Leningrad. 

5773 108 



^H 



40 cm high, sometimes bearing one scalelike leaf; inflorescence branching, 
rounded -paniculate, compressed, many -flowered; pedicels 0.5—1 cm long; 
calyx campanulate, parted to Y 3 — / 2 , red, glabrous, the sepals orbicular or 
acuminate; petals red (violet in herbarium), oblong, tapering into long 
claw, rounded at the apex, to 2 cm long, 3 times as long as calyx; stamens 
and pistils shorter than petals, with 2 erect styles. July. 

Rocky taluses and forest slopes.— Far East: Uss. Endemic. Described 
from Ternei Bay on the Pacific coast. Type in Leningrad. 



Genus 706. SAXIFRAGA * .L. 

L.Gen.pl.ed.l (1737) 131; Sp.pl.ed.l (1753) 398; Engl.et Irmsch.in Pflnzr.H. 67, 117,1V, 1 (1916) 1. 

Calyx 5 -sect, gamophyllous; petals 5; stamen 10; pistils with 2 free 
styles; ovary 2 -locular, nearly free or half-sunken; fruit a capsule 
dehiscing along inner suture between styles, with persistent styles, 
and hence bicorn; seeds small, round, oblong or fusiform. Herbaceous 
perennials or annuals, with rosette of radical leaves and scapiform stem 
or with leafy perennial or dying -off shoots and annual flowering stem; 
leaves variable in shape and size, often spatulate usually pubescent or 
ciliate, sometimes glabrous. 

1. Leaves opposite; cauline shoots densely leafy, cespitose; flowers 
solitary, subsessile, pink or violet 77. S. oppositifolia L. 

+ Leaves alternate 2. 

2. Leaves smooth-margined, dentate, ciliate, glandular or cartilaginous, 
but never secreting lime on the margin and without marginal 

pits 3. 

+ Leaves smooth-margined or dentate, ciliate or glandular, always with 
lime -secreting pits on the margin, or else lime secreted throughout 
the margin 6 3. 

3. Flowers actinomorphic 4. 

+ Flowers zygomorphic; petals varying in shape and size (Section 

Diptera Borkh.) 62. 

4. Stem leafless or 1 - or 2-bracted; leaves in radical rosette. 

(Section Boraphylla Engl.) 5. 

+ Stem leafy 27. 

5. Plants loosely cespitose; flowers to 15 cm in diameter; petals white, 
rounded, long -clawed; anthers black 23. S. merkii Fisch. 

+ Plants not cespitose; flowers smaller and not as above 6. 

6. Leaves orbicular to reniform 7. 

+ Leaves oblong, oblong -oval, oval -rhomboid, usually with cuneate base 

and tapering 14. 

7. Petioles with dry, membranous, auriculate stipules 

1 . S. nudicaulis D. Don. 

+ Petioles exstipulate 8. 

8. Petals lobate -incised 9. 

+ Leaves crenate -dentate, but not lobate; petals white, white with red 

dots, or red 10. 



From the Latin sa x u m, rock, and fra ng o, I break. 



109 



9. Leaf lobes 7—9, obovate; petals white, with 2 confluent yellow 

spots 1. S. sieversiana Sternb. 

+ Leaf lobes rhomboid -elliptic, tapering downward, truncate above; 

petals white, without spots 3. S. korshinskyi Kom. 

10. Leaves delicate, slender, and somewhat fleshy, more or less 

large 11. 

+ Leaves compact, coriaceous, small 13. 

11. Inflorescence sparse, spreading; sepals oblong-oval; anthers 
reniform 12. 

+ Inflorescence compressed, subcapitate; sepals elongate or linear, 

anthers oblong 8. S. manshuriensis (Engl.) Kom. 

12. Leaf teeth oval -deltoid, gradually tapering into mucro 

4. S. reniformis Ohwi. 

+ Leaf teeth oval, short -acuminate 5. S. punctata L. 

13. Petals white or pinkish 6. S. nelsoniana D. Don. 

+ Petals vinaceous 7. S. purpurascens Kom. 

14. Leaves cuneate, rounded at the apex, pectinate -dentate 15. 

+ Petals oblong, ovate, or rounded -rhomboid, with rounded or cuneate 

base, dentate throughout margin or entire 19. 

15. Leaves with auriculate stipules at base 16. 

+ Leaves without stipules at base or, if stipules present, then without 

auricles and rounded 17. 

Leaves sharply cuneate, densely glandular throughout blade and 

petiole 10. S. redowskiana Sternb. 

+ Leaves reniform -cuneate, glabrous or sparsely hairy on the margin 

9. S. astilbeoides A. Los. 

17. Inflorescence compoundly branching, corymbiform-paniculate .... 18. 
+ Inflorescence simple, racemose 13. S. calycina Sternb. 

18. Petals to 3.5 mm long 12. S. grand ipetala (Engl.) A. Los. 

+ Petals to 2 mm long 1 . S. dahurica Willd. 

19. Leaves subsessile 20. 

+ Leaves with conspicuous petioles 22. 

20. All flowers normally developed 21. 

+ Only the terminal flower developed, all others transformed into 

compact leaf buds 20. S. foliolosa R. Br. 

21. Sepals obtuse 21. S. redowskyi Adams. 

+ Sepals acute 22. S. stellaris L. 

22. Plants to 50 cm high; stem robust 23. 

+ Plants to 25 cm high; stem slender 24. 

23. Petals white; leaves ovate, serrate -dentate 

18. S. sachalinensis F. Schmidt. 

+ Petals reddish or greenish; leaves crenate -sinuate 

19. S. hieraciifolia Waldst. 

24. Petioles as long as or shorter than leaf blade; inflorescence 
compressed, paniculate, few-flowered 25. 

+ Petiole longer than the blade; inflorescence corymbiform-paniculate, 
many -flowered 26. 

25. Entire plant slightly reddish; petals 1.5 mm long 

15. S. tenuis (Wahlenb.) Sm. 

+ Entire plant green; petals 2.5—3 mm long 14. S. nivalis L. 



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26. Stem reddish violet; calyx, pedicels, and stamens black violet 

17. S. melaleuca Fisch. 

+ Stems light green; sepals, pedicels, pistils and stamens black- 
violet 16. S.tilingiana Rgl. 

27. Plants with underground rhizome, not giving rise to leafy cauline 
shoots, or else annual plants 28. 

+ Plants with developed above-ground leafy shoots, cespitose 45. 

28. Leaves entire, ciliate, glandular or hairy. (Section H i r i c u lu s 

(Haw.) Tausch 29. 

+ Leaves lobate, dentate or dissected 33. 

2 9. Plants with long filiform shoots, terminating in rooting rosettes; 

leaves ciliate or glandular 31. 

+ Plants without filiform shoots 30. 

30. Plants without radical leaves; pubescence not rufous 

24. S. parnassioides Rgl. et Schmalh. 

+ Plants with a radical rosette; pubescence rufous. . . . 25. S. hirculus L. 

31. Leaves glandular on the margin; calyx with subtruncate base; sepals 
glandular on the margin 27. S. komarovi A. Los. 

+ Leaves ciliate on the margin. Plants green; calyx with infundibular 

or rounded base 32. 

32. Calyx dissected to base, with rounded base, rarely glandular; flowers 
gaping, as long as pedicels 26. S. flagellaris Willd. 

+ Calyx dissected to the middle, densely glandular, with infundibular 

base; flowers with very short pedicels, campanulate 

2 8. S. setigera Pursch. 

33. Leaves orbicular, emarginate, dentate or spatulate 34. 

+ Leaves oblong, dentate only in upper part. (Section T r i d a c ty 1 it e s 

Griseb.) 44. 

34. Seeds rounded. Annual plants with decumbent, ascending, less often 
erect shoots. (Section Cymbalaria Griseb.) 35. 

+ Seeds oblong. Perennial plants; stems always erect 36. 

35. Leaves with 5—7 broadly oval lobes, with light brown veins below .... 
31 . S. huetiana Boiss. 

+ Leaves with 5—13 large, triangular -oval teeth, without light brown 

veins 32. S. cymbalaria L. 

36. Plants with radical bulbils; leaves thin, glabrous or pubescent; ovary 
semi -imbedded. (Section N a p h r o p hy 1 1 um Gaud.) 37. 

+ Plants without radical bulbils; leaves coriaceous; ovary free. 

(Section Miscopetalum Haw. ) 

. 30. S. coriifolia (Somm. et Lev.) Grossh. 

37. Inflorescence crowded, subcapitate; flowers minute -pediceled, 
surrounded by approximate bracts 41. S. bracteata D.Don. 

+ Inflorescence corymbiform -paniculate or flower solitary in leaf axils 
or at tips of branches 38. 

38. Inflorescence corymbiform -paniculate, many -flowered; leaves 
pubescent 39. 

+ Flowers solitary, few; leaves glabrous 40. 

39. Stems numerous; radical rosette dense. Soft -hairy plants; leaves 
with twice 3— 4 -sect oblong lobes 35. S. irrigua M. B. 

+ Stems solitary. Rugose -hairy plants; leaves with 5—9 rounded, 

subtruncate lobes 36. S. granulata L. 

40. All flowers normally developed 41. 



Ill 



+ Only terminal flower developed, the others transformed into leaf 

buds-bulbils 37. S. cernua L 

41. Radical leaves with 3—5 oblong, acute lobes with cuneate or rounded 
base (Arctic forms) 42. 

+ Radical leaves with 5 — 9 broadly ovate or triangular -ovate acuminate 

lobes 43. 

42. Petals 3—4 mm long 40. S. rivularis L. 

+ Petals 10— 12 mm long 42. S. exilis Stephan. 

43. Petals 5 — 7-nerved, to 14 mm long 38. S. sibirica L. 

+ Petals 3 -nerved, to 20 mm long 39. S. mollis Schmidt. 

44. Annual plants; pedicels twice as long as petals; petals twice as long 

as sepals 33. S. tridactylites L. 

+ Biennial plants; pedicels 3—4 times as long as flowers; petals 3—4 

times as long as sepals 34. S. adscendens L. 

45. Leaves spiny-ciliate or glandular, mucronate. (Section Trachy - 
phyllum Gaud.) , 46. 

+ Leaves unarmed, lobate or dissected or entire, muticous 51. 

46. Tufts spherical; flowers solitary, subsessile; leaves rounded at the 

apex, concave, long-spiny-ciliate, straw-colored or silvery 

58. S. escholtzii Sternb. 

+ Tufts of indeterminate shape; inflorescence many-flowered 47. 

47. Upper leaves at ends of shoots involute, forming coils 48. 

+ Leaves at ends of shoots not forming coils 49. 

48. Leaves strongly concave, 24 mm long 57. S. cherlerioides D.Don. 

+ Leaves flatter, lustrous on the outside, 6— 8 mm long 

56. S. firma Litw. 

49. Leaves keeled below; petals 5— 7 mm long 50. 

+ Leaves slightly concave below; petals less than 5 mm long 

55. S. spinulosa Adams. 

50. Leaves pectinate-ciliate 54. S. bronchialis L. 

+ Leaves glandular-ciliate on the margin .... 53. S. anadyrensis A. Los. 

51. Leaves entire, oblong-oval or linear, thickish, glabrous, ciliate- 
margined, convex below, with a pit above, not densely covering shoots 
(Section Xa nt hi s i o n Griseb.) 59. S. aizoides L. 

+ Leaves not as above, without pit. (Section Da cty lo id e s Tausch.) . . 
52. 

52. Leaves entire 53. 

+ Leaves dissected or lobate 54. 

53. Leaves cartilaginous on the margin, glabrous; flowers more than 1 cm 
in diameter; petals yellow, 5 -nerved 29. S. serpyllifolia Pursch. 

+ Leaves without cartilaginous margin, glandular; flowers 0.5—0.6 cm 

in diameter; petals white, 3 -nerved 43. S. androsacea L. 

54. Flowers solitary, less often 1 or 3 55. 

+ Flowers numerous in inflorescence 5 7. 

55. Calyx constricted at base of sepals; flowers 1 cm in diameter 

, 44. S. sileniflora Sternb. 

+ Calyx not constricted; flowers 0.4— 0.7 mm in diameter 56. 

56. Petals 4.5— 5 mm long, twice as long as sepals 

47. S. terektensis Bge. 



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!■■■■■■■■■ 



+ Petals 5 — 7mm long; 3 limes as long as calyx 46. S. caespitosa L. 

57. Petals 3 times as long as sepals, white 45. S. lactea Turcz. 

+ Petals not more than twice as long as sepals 58. 

58. Petals twice as long as sepals, yellow or while; Leaves with 3 
prominent veins petiolatc 59. 

+ Petals white, yellow -green, sometimes reddish, slightly Longer than 

sepals; leaves without prominent veins (i,) - 

59. Densely cespitose plants; Leaves densely overlapping, to L2 mm Long 
51. S. adenophora K. Koch. 

+ Loosely cespitose plants; Leaves not densely overlapping, to 20 mm 

long 50. S. exarata VMI. 

GO. Petals white, orbicular or rounded oval; leaves verticillate 

52. S. verticillata A. Los. 

, .. + Petals greenish or reddish; leaves approximate 61. 

1 44 

61. Cauline shoots columnar, densely imbricate leaved ; 

4 9. S.pontica A I bow. 

+ Cauline shoots not columnar, loosely leaved 48. S. moschata Wull'. 

62. Leaves rounded -reniform 78. S. cortusifolia Sieb. et Zucc. 

+ Leaves oblong or ova) 79. S. oblongifolia Nakai. 

63. Leaves with several lime -secret Lng pits on the margin (Sed Lon 
Kabschia Engl.) 65. 

+ Leaves secreting lime throughout margin; pits absent. (Section 

Euaizoonia (Schott) Engl.) 64. 

64. Leaves gradually tapering toward apex; petals purple 

61 . S. kolenatiana Rgl. 

+ Leaves abruptly narrowed at apex; petals white or pinkish 

60. S. cartilaginea Willd. 

65. Flowers solitary (,,i - 

+ Flowers in a panicle or corymb f >8. 

66. Flowers purple; leaves obi use, s ma 1 1 , densely eoverifig slender, 
columnar shoots 75. S. columnaris Schmalh. 

+ Flowers white or yellow; leaves acuminate, less densely covering 

the stems ( '7. 

67. Leaves with 5 pits, approximate, with reflexed tips 

76. S. dinnikii Schmalh. 

+ Leaves with one apical pit at verticillate, erect or reflexed 

72. S. carinata ( Jetting. 

68. Leaves obtuse or with apical thickening 69. 

+ Leaves acuminate 70. 

69. Leaves with 3-5 pits; petals white .... 74. S. albertii Rgl. et Schmalh. 
+ Leaves with 7-9 pits; petals yellow 73. S. kotschyi Boiss. 

70. Flowering stems glabrous 71 . 

+ Flowering stems pubescent 74. 

71 . Cauline shoots with imbricate leaves 72. 

+ Cauline shoots with verticillate leaves 73. 

72. Leaves with one, often inconspicuous pit, rarely with 3 5 pits; petals 

slightly longer than sepals, 1— 3-nerved 

69. S. caucasica Somm.^el Lev. 

+ Leaves with 5—7 pits; petals white, more than twice as Long as sepals, 
145 5 -nerved 71. S. kuznezowiana Oett. 



13 



73. Verticillate leaves erect, appressed to shoot. . . . 64. S. colchica Albow. 
+ Verticillate leaves declinate or divergent 

63. S. subverticillata Boiss. 

74. Leaves flat or slightly keeled, grooveless on upper side 75. 

+ Leaves keeled or convex below, with groove on upper side 77. 

75. Petals 1 -nerved; leaves reflexed .... 67. S. scleropoda Somm. et Lev. 
+ Petals 3— 5 -nerved; leaves horizontally spreading or erect 76. 

76. Leaves lustrous, erect, conspicuously veined, to 20 mm long and 4 mm 

broad; petals 5 -nerved, conspicuously clawed 

66. S. pseudolaevis Oett. 

+ Leaves dull, concave, about half as long as in preceding species; 

petals 3 -nerved, without claw 65. S. laevis M. B. 

77. Leaves keeled below, tapering at the apex, spiny, long-lanceolate or 
oblong-linear 62. S. juniperifolia Adams. 

+ Leaves concave below, oblong-lanceolate or oblong -oval 78. 

78. Flowering stems covered with rather long, white hairs 

70. S. desulavyi Oett. 

+ Flowering stems covered with small rufous hairs 

68. S. abchasica Oett. 



Section 1 . BORAPHYLLA Engl., Ind. cult, in Verh. Zool. -bot. Ges. Wien 
(1869) 521.- Micranthes Haw., Syn. pi. succ. (1812) 320; Enum. Sax. 
(l 821 ) 45 (pro gen.).— Calyx with flat base; ovary superior, not sunken; 
filaments usually fusiformly dilated; capsule slightly dissected, with short 
style; seeds oblong. Flowering stem leafless; leaves in radical rosette, 
usually dentate, incised or lobate. 



Cycle 1. Nudicaules A. Los. — Grex P u nc t at a e Engl, et Irmsch., 
1. c, 6, p. p. — Small, slender plants; stems glabrous. 

1. S. nudicaulis D.Don, Transact. Linn. Soc. XIII (l82l) 366; Engl, et 
Irmsch., 1. c.,15.— S. neglecta Bray in Sternb., Suppl. I (1822) 9. — 
S. vaginalis Turcz. in Ldb. Fl. Ross. II (l 844) 820. - Ic. : Sternb., 1. c, 
tab. 6. 

Perennial, subglabrous, small; rhizome oblique; stems numerous, erect, 
slender, minutely sulcate, 5—18 cm high, with small trilobate bracts at base of 
inflorescence; radical leaves reniform, dentate (lobate), the teeth oval, 
acute, to 2 mm long, the blade to 2 cm broad, 1 .5 cm long, the petiole longer 
than blade, with membranous, broad, auriculate stipules; inflorescence 
broadly paniculate -corymbiform; pedicels filiform, slightly longer 
than flowers; calyx with triangular and ovate sepals, 1.5 mm long, dissected 
to base; petals with conspicuous claw, 2 — 3 times as long as calyx, yellowish; 
stamens with dark anthers, exceeding corolla; pistils with short style. 
July. (Plate VIII, Figure 9). 

Moist riverbanks, mountains.— Arctic: Arc. Sib., Chuk., An.; E. Siberia: 
Lena-Kol.; Far East: Ze.-Bu., Okh. ; Gen. distr. : Ber. Described from 
N. America. Type in Banks' herbarium in America. 

Note. F. st olo ni f e r a Kjelmann (ex Engl, et Irmsch., 1. c, 16), with 
decumbent leafy stems and scattered leaves, occurs in Sakhalin. 



114 



2. S. sieversiana Sternb., Suppl. II (1831) 16; Engl., Mon. Sax. (1872) 
140; Engl, et Irmsch.,1. c.17.— S. sibirica Sternb., Rev. (l 810) 23, 

non L. — S. reticulata Cham, in Linnaea VI (1831 ) 555.— S.bimaculata 
Turcz.in Bull. (Soc. Nat. Mosc. (1840 ) 71; Ldb., Fl. Ross. II, 220. - 
Micranthes sieversiana tSternb.) Kom., Fl. Kamtsch. II (192 9) 
212. - Ic: Sternb., Suppl. II (1831 ) ab.25, f.2. 

Perennial; stem slender, delicate, slightly branching, 10—15 cm high, 
with one leaf at base of fork, glabrous in upper part, glandular below; 
leaves rounded -reniform, 1 .5 cm broad, 1 cm long, hairy on both sides, 
with 7 or 8 obovate, obtuse lobes; petioles densely pubescent, twice as 
long as blade; flowers 1 or 2 at ends of stems and branches; bracts 
oblong, acute, 3—5 cm long; pedicels slender, 0.5—3 cm long; calyx with 
divergent, oblong or oblong -triangular sepals, 1.5 mm long; petals white, 
with 2 confluent yellow spots, broadly oval, tapering upward, obtuse, short - 
clawed, 4 mm long; stamens shorter than petals; pistils with oval ovary 
and short divergent styles. 

Far East: Okh. (Stanovoi Range). Endemic. Described from the vicinity 
of Okhotsk. 

Note. There are no ecological or biological data in the literature 
or on labels of the very limited herbarium material. 

3. S. korschinskyi Kom.in A.H. P. XVIII (1900) 436; Fl. Mansh. II 
(1903) 416; Engl, et Irmsch., 1. c. 17. - S. sp. Korshinsky in A.H. P. XXII 
(1892) 341, No. 240.- Ic: Kom., Opred. rast. Dal'nevost. kraya I (1930) 
tab. 182, f. 5-6. 

Perennial; stem to 18 cm high, slender, delicate, glabrous; petioles 
slender, 1— 4 cm long, leaf blade, 3— 5 -fid, the middle 3-lobed,the lateral 
bi-lobulate, all lobules oval -elliptic, obtuse, tapering downward, 6— 7 mm 
long, 2—4 mm broad; leaves covered above with small epidermal papillae, 
smooth below, often reddish; flowers (2—6) in loose inflorescence; 
pedicels filiform, 1 — 1.5 cm long, with small, linear, acute bracts; sepals 
oblong, 1 mm long; petals white; capsule dehiscing at the very apex. 
June -July. (Plate VIII, Figure 10.). 

Growing in tufts on moist rocks, slopes, among mosses, forming with 
mosses a thin, easily removable cover.— Far East: Ze.-Bu. (basin of the 
Tyrma and Bureya rivers). Endemic. Described from the village of 
Kulakovka on the Bureya River. Type in Leningrad. 



Cycle 2. Punctatae (Engl, et Irmsch.) A. Los. — Grex P u nc t at a e 
Engl, et Irmsch., 1. c, 6, p. p. — Rather large plants; leaves coarsely 
dentate, reniform; stems pubescent; styles rather long. 

4. S. reniformis Ohwi in Acta Phytotaxonom. et geobot. II (1933), No. 125.- 
S. aestivalis Fr. Schm., Reise Am. u. Sach. (1868) 133.— S. punctata 
Hult., Fl.Kamtsh. Ill (1929) 25, p.p.- S. punctata f. corymbosa Engl, 
et Irmisch., 1. c, 1 1 . 

Perennial; rhizome creeping, thick; stem erect, 10—30 cm high, glabrous 
below and whitish-pubescent — like the pedicels — above; petioles 5—10 cm 
long, sulcate above, leaf blade fleshy, reniform, with cordate base and large 



115 



oval-deltoid teeth; inflorescence branches and pedicels rather thick; 
flowers 0.6—0.7 cm in diameter; calyx with divergent, oblong, glabrous, 
obtuse sepals 1.5 mm long; petals oblong -oval, white, refiexed, 3 mm long, 
obtuse, with small claws, 1 -nerved; filaments dilated at base; anthers 
reniform; styles short, thick. July. (Plate VIII, Figure 7). 

Along mountain streams.— Far East: Sakh. Gen. distr.: Japan. 
Described from Mt. Tosso in Sakhalin. Type in Tokyo. 

Note. Distinguished from S. p u n c t a t a L. by its short inflorescence 
branches and its narrower and gradually tapering leaf teeth. 

5. S. punctata L., Sp.pl. ed. I (1753) 401; Ldb., Fl. Ross. II, 215; Engl, 
et Irmsch.,1. c, 9; Kryl., Fl. Zap. Sib. VI (l 931 ) 1424; Kom., Fl. Kamch. II 
(1929) 209.- S. aestivalis Small, N. Amer. Fl. XXII, 2 (1905) 147. 

Perennial; rhizome creeping; stem leafless, erect, 10—50 cm high, 
pubescent in upper part, glandular -hairy in inflorescence; petioles 
long, pubescent, 2—4 times as long as blade; leaf blade puberulent on 
both sides or glabrous, rounded -reniform, broadly cordate at base, 1.5— 8 cm 
long, 2— 9 cm broad, dentate, with short -acuminate teeth; inflorescence a 
many-flowered, spreading or congested panicle; flowers small; calyx 
glabrous, dissected to 2 / 3 , with ovate, acuminate teeth, refiexed after 
anthesis; petals white with orange dots or slightly pinkish, obovate or 
oblong, short-clawed, 3— 3.5 mm long, twice as long as calyx; filaments 
fusiformly dilated; anthers orange, reniform; capsule oblong, June — July. 
(Plate VIII, Figure 6). 

Arctic and polar regions, moss and moss -and -lichen marshy tundra, near 
snow, stream and river banks, less often alpine and subalpine meadows. — 
Arctic: Arc. Sib., Chuk., An.; European part : Dv. -Pech., V. -Kama 
(N.Urals); W.Siberia: U. Tob., Ob, Alt.; E.Siberia: Yenis., Ang.rSay., 
Lena-Kol.; Far East.: Kamch., Sakh., Ze. -Bu., Uss.; Central Asia: 
Dzu. -Tarb. Gen. distr.: Mong., N. Am. Described from Siberia. Type in 
London. 

Note. Inflorescence varies greatly from broadly paniculate to narrowly 
ovoid; Arctic and bald -mountain forms are characterized by their 
larger corolla, thicker pedicels, and more compressed inflorescence. 
Consistency and pubescence of leaves also vary. Transitional forms 
between S. punctata L. and S. nelsoniana D. Don may be observed. 

6. S. nelsoniana D.Don in Trans. Linn. Soc. XIII (1821) 355; Kom., Fl. 
Kamch. II (192 9) 209.— S. punctata var. krauseana Engl, ex Kurtz in 
Engl., Bot. Jahrb.XIX (1895) 462 9.- S. punctata var. nelsoniana 
(D.Don) Engl., Mon. Gatt. Sax. (1872) 139.- Micranthes nelsoniana 
Small, N. Amer. Fl. XXII, 2 (l 905) 140. - S. p u n c t at a B. Fedtsch., II. Com. 
(l 905) 62. — S. p u nc t a t a var.typica f.coarctata Engl, et Irmsch., 
I.e., 11. 

Perennial; rhizome slender, with underground shoots; stem leafless, 
erect, 10—30 cm high, sparsely hairy, glandular; leaves in rosette 5 or 6, 
firm, slightly fleshy, bright green, paler below, glabrous or puberulent, 
rounded -reniform, with deeply cordate base, with acuminate, large, oval, 
glandiferous teeth, 1—3 cm long, 1—4 cm broad; petioles slightly longer than 
blade, reddish; inflorescence ovoid, compressed; pedicels slender, densely 
glandular, with small linear bracts at base, the lowest bracts often dentate; 



116 



flowers 0.6—1 cm in diameter; sepals oblong, purple; petals oblong or 
oval, white or pinkish, twice as long as sepals, tapering to short claw; 
styles divergent. July. 

Rocks, pebbles, alpine meadows, mountain stream banks. — Arctic: 
Arc. Sib., Chuk., An.; Far East: Kamch. Gen. distr. : Ber., N. Am., 
Described from the vicinity of Petropavlovsk-Kamchatskii. 

Note. Distinguished from S. p u nc t at a L. by its size, compressed 
inflorescence, fleshy, mucronate -toothed leaves. 

7. S. purpurascens Kom. in Fedde, Rep. sp. nov. XIII (1914) 167; 

Fl. Kamch. II (1929) 211. — S. p u nc t a t a var.typica f . p u r p u r a s c e n s 
Engl, et Irmsch., 1. c, 10. — Micranthes purpurascens Kom., Fl. 
Kamch. II (1929) 211. 

Perennial; rhizome thickened, branching, oblique, 5 — 10 cm long; stem 
6—32 cm high, scattered -pubescent; radical leaves long-petioled, cordate- 
rounded or cordate-reniform, less often hairy, 3—6 cm in diameter, 
with equal, large, rounded -acuminate teeth, pubescent at the margin, green 
or reddish; petioles longer than blade, reddish, pubescent with white hairs, 
like the stem; panicle not dense, broadly oval in outline, individual 
peduncles to 2 cm long, glabrous or minutely glandular; bracts at base 
of peduncles dentate, linear; pedicels 4 — 12 mm long, filiform; sepals 
1.5— 2 mm long, reflexed, black-purple; petals narrowly oblong-oboval, 
black-purple, twice as long as sepals; capsule ovoid, with divergent styles. 
August. (Plate VIII, Figure 8). 

Alpine zone: alpine meadows, rock crevices in rock streams.— Far East: 
Kamch. Endemic. Described from the Mt. Trub in Kamchatka. Type in 
Leningrad. 

8. S. manshuriensis (Engl.) Kom. Fl. Mansh. II (1903) 415.- S. punctata 
var. ma ns hu r i e ns i s Engl. Mon. Gatt. Sax. (l 872) 139; Engl, et Irmsch., 

1. c, 7. 

Perennial; stems 15— 30 cm high, strong, densely glandular -hairy, sometimes 
with 1 cauline leaf; leaves long-petioled, sparsely pubescent, sometimes as 
long as stem; the blade 4—7 cm long, 6—8 cm broad, reniform, fleshy, glabrous 
or sparsely hairy, with large, broad, mucronate teeth; inflorescence a 
many -flowered, compressed, ovoid or spherical panicle, with lanceolate, often 
3 -fid bracts at base of peduncles and linear bracts at base of slender, to 
1.5 cm long, glandular pedicels; sepals linear, subobtuse, 2— 2.5 mm long, 
0.5 mm broad, pubescent; petals narrowly oblong, white, obtuse, 5—5.5 mm 
long; filaments as long as petals; anthers narrowly claviform, attaining 
6 mm after anthesis; pistils with oblong ovary and with style, as long as 
stamens. August. (Plate VIII, Figure 5). 

Riverbanks, moist soils, among bushes, in dense grass, less often on 
stony slopes near streams.— Far East:Uss. Gen. distr.: Jap.-Ch. Described 
from Manchuria. Type in London; cotype in Berlin. 



Cycle 3. Dahuricae Engl, et Irmsch., 1. c, 1 9. — Leaves cuneate, 
orbicular or rounded -rhomboid above, with large teeth; petals clawed. 



117 



9. S. astilbeoides A, Los. sp. nova in Addenda VIII, p. 3 72. — S. dahurica 
Kom. and Alis., Opred. rast. Dal'nevost. Kraya I (l 931 ) 608, non Willd. — 

Perennial; stem erect, ribbed, sometimes compressed, branching, 
minutely glandular, to 25 cm high; leaves light green, red below, reniform- 
cuneate, with large, oval mucronate teeth in upper half, glabrous or with 
sparse hairs at the margin, to 2.5 cm long, 3 cm broad; petioles sulcate, 
canaliculate above, minutely glandular, to 8 cm long, with auriculate acute 
stipules at base; inflorescence paniculate, spreading, widely branching, the 
lower branches arising from middle of stem, somewhat arched-upcurved, 
bearing at the base 1 bract, replaced in low-growing specimens by narrowly 
lanceolate bracts; inflorescence branches profusely branching, terminating 
in straight branchlets.each bearing 4 — 8 flowers on slender, glandular, 
slightly curved pedicels as long as branchlets (5 mm), with small bracteoles; 
flowers 0.5—0.6 cm in diameter; calyx glabrous, reddish, dissected to the 
middle, with triangular sepals; petals white, oval, to 2.5 cm long, tapering 
to claw; stamen as long as petals, with slender filaments and black -violet 
anthers; ovary broadly oval, with curved styles. July— August. (Plate VIII, 
Figure 4). 

Stony taluses, pebbly slopes.— Far East: Uss. Endemic. Described 
from the Botcha River. Type in Leningrad. 

10. S. redowskiana Sternb., Rev. Sax. Suppl. II (1831 ) 44.— S. dahurica 
Ldb., Fl. Ross. II, 212, p. p., non Willd. — S. dahurica f. willdenowii 
Engl, et Irmsch., 1. c, 21 p. p. — S. dahurica j3 latifolia Sternb., 
Suppl. I (1820) tab. 5. — Ic. : Engl, et Irmsch., 1. c. (sub S. dahur i c a) 

f. 4 H. M. - 

Perennial; rhizome very slender; stems usually solitary, to 20 cm 
high, sulcate, densely and minutely glandular; leaves firm, thickish, 
light green, rounded -rhomboid, abruptly narrowing to long slender petiole, 
incised in upper half into oval -triangular mucronulate teeth; the blade 
to 2.5 cm long and broad, covered throughout — like the petioles — with 
dense green glands borne on basally broadened stalks; petioles to 6 cm 
long, broadening downward and bearing acuminate scarious, auriculate 
stipules; inflorescence dense, branching, corymbiform, their oblique 
ascending, densely glandular peduncles with large oblong -lanceolate 
bracts; flowers to 7 mm in diameter; sepals narrowly triangular, acute, 
often purple, glandular and glabrous; petals to 2.5 mm long, about twice 
as long as sepals, white, yellowish or pinkish, oblong-oval, with long claw; 
stamens shorter than petals; filaments slender, slightly dilated toward 
base, anthers violet; capsule ovoid -triangular, dehiscing to the middle; 
stigmas subsessile. July — August. (Plate VIII, Figure 3). 

Stony taluses, among stones, on rocks, and balds. Arctic: Arc. Sib., An.; 
E.Siberia: Lena-Kol.; Far East : Ze. -Bu., Uda, Okh. Endemic. Described 
from the Yablonovyi Range. 

Note. Mention should be made of a many -stemmed form, observed 
on seashores (Taui Bay, mouths of the Ayan and Anadyr rivers), lower - 
growing, often with reddish leaves, branching cauline shoots, numerous 
leaf rosettes, and several stems. 



118 



11. S.dahurica Willd., Sp.pl. II, 1 (1799)645; Ldb., Fl. Ross. I, 212 p. p., 
Engl, et Irmsch., I.e., 21 p.p. 

Perennial; rhizome vertical, short; stem erect, leafless, glabrous or 
sparsely glandular, ribbed, 5-10 cm high; all leaves radical, light green, 
oval or rounded, incised in upper part into large, narrowly triangular, acute 
teeth (5—8), tapering to long, slender petioles, broadening downward, with 
a thin, membranous border below; leaf blade glabrous, sparsely glandular 
on the margin, 1.5—2 cm long and broad; inflorescence corymbiform- 
paniculate, branching, the branches and pedicels minutely glandular; 
bracts small, lanceolate or linear; calyx dark purple, glabrous, with 
rounded base and triangular, acute sepals; petals white or pink, oblong- 
lanceolate, with long claw, to 20 mm long, 0.5 mm broad; filaments white, 
slightly dilated toward base; anthers purple, spheroid, the stamens about 
as long as petals; ovary rounded -oval; stigmas subsessile. June — August. 
(Plate VIII, Figure 1 ). 

Balds, rock streams.— E.Siberia: Dau., Ang. -Say., Lena -Kol. (S. part 
of the Dzhugdzhur Range). Endemic. Described from Dauria. Type in 
Berlin. 

12. S. grandipetala (Engl, et Irmsch. ) A. Los. comb. nova. — S. d a hur i c a 
f. g r a nd i p et a la Engl, et Irmsch., 1. c, 22. 

Perennial, representing the Arctic race of S. dahurica Willd., 
distinguished by the larger petals to 3.5 mm long and 2 mm broad, 
sometimes also by broad membranous rounded stipules without auricles 
(Krause's specimens from Litke Bay, cited by Engler). Species requiring 
verification, with very little data available. July— August. 

Riverbanks pebbles.— Arctic: Chuk., An. Endemic. Described from the 
Chukchi Peninsula, from Litke Bay. Type in Berlin, cotype in Leningrad. 

13. S. calycina Sternb., Rev. Sax. Suppl. II (l 831 ) 10.- S.dahurica 
f.calycina Engl, et Irmsch., 1. c, 22. — S. d a hu r i c a Ldb., Fl. Ross. II, 
1, 212, non Willd.; Kom., Fl. Kamch. Il(l93l)264.-S.unalaschkensis 
Hulten, Fl. Kamtsh. Ill (1929)30.- Ic. : Sternb., 1. c, tab. XXI. 

Perennial; rhizome elongated, oblique; stem erect, sulcate, sparsely 
glandular, 5—10 cm long; leaves light green, oval- rhombic -cuneate, 
tapering to broad petiole, deeply incised in upper half into acute, erect 
teeth; the blade together with petiole to 5 cm long, 2 cm broad, glabrous, 
covered on the margin — especially in lower part — with long, soft, white 
hairs; stipules absent; inflorescence racemose, not branching, ovoid; 
pedicels to 15 mm long, densely glandular and hairy, with linear -lanceolate 
bracts appressed and slightly shorter than pedicels; flowers to 1 cm in 
diameter; sepals oval-oblong, acuminate, 2.5 mm long, often purple, reflexed 
at anthesis; petals yellowish white, oblong-oval, clawed, 1 -nerved, 3 mm long; 
stamens half as long as petals, with bright yellow fusiform filaments and 
minute brown anthers; ovary oblong, with short styles; capsule oblong, 
dehiscing nearly to the middle. June — August. (Plate VIII, Figure 2). 

Arctic and alpine zones, gravels, dry lichen carpets, taluses. — Arctic: 
Arc. Sib., Chuk., An. ; Far East: Kamch. Endemic. Described from the 
shores of Lavrentiya Bay (after Chamisso's specimen). Type in Berlin. 



119 



(153) 




PLATE Vm. 1-Saxifraga dahurica Willd • 2 Sn.i ■ 
b - S. punctata L i».f- -7 - _ . _ -case, &-s 



.leaf; 7 - S.reniformi 



a Sternb. 
manshuriensis (Engl.) Kom.,leaf; 



S.nudicaulis D.Don., a) petiole ba e/i - s kor'sh ^'"P'"""" to.Ie.fi 9- 

S.korshinskyiKom. > a)leaf; 11 _ s . fnl ,- n1 „.. 



11 - S.foliolosa R.Br. 



120 



Note. There are forms with branching inflorescences of a type 
transitional between S.calycina Sternb. and S. d a h u r i c a Willd. 



Cycle 4. Nivales A. Los. — Gres Nivali -virginiensis Engl, et 
Irmsch., 1. c, 22 p. p. — Inflorescence capitate, few-flowered; petiole as 
long as the blade. 

14. S. nivalis L., Sp.pl. ed. I (1753)401; Ldb., Fl. Ross. II, 21 3; Engl, et 
Irmsch., 1. c, 29; Kom., Fl. Kamch. II (1929) 212; Kryl., Fl. Zap. Sib. VI, 
1423.- S.pauciflora Sternb., Suppl. I (1822) 6, tab. 4.- Micranthes 
nivalis Small, N.Amer. Fl. XXII, 2 (1905) 136.- Derma sea nivalis 
Haw., Enum. Sax. (1821) 9. 

Perennial; stem solitary, leafless, 5 — 15 cm, less often to 20 cm high, 
covered like the pedicels with remote glandular hairs, especially in 
upper part, rosette leaves obovate or ovate, 0.6—3 cm long, 0.3—2 cm broad, 
cuneately tapering to broad petiole as long as the blade; blade glabrous, 
firm, obtusely dentate -margined in upper part, hairy -margined in lower 
part; flowers few, in a compressed capitate, less often loose corymbiform- 
paniculate inflorescence; bracts oblong or linear; pedicels short, glandular; 
255 calyx incised to the middle, with bluntly acuminate, erect sepals; petals 
white, elliptic, subobtuse, tapering to short claw, the petal together with 
the claw to 3 mm long and 1.5 mm broad, slightly longer than calyx; capsule 
ovoid, deeply dehiscing, 7 mm long, 5 mm broad. July —August. 

Alpine and arctic regions, tundra, rock crevices, stony and shady river - 
banks.— Arctic: Arc. Eur., Nov. Z., Arc. Sib., Chuk., An.; European part. 
Kar. -Lap., Dv.-Pech., V. -Kama (N. part); W.Siberia: Ob (N. part), Alt. 
(rarely); E.Siberia: Yenis. (N. part), Lena-Kol., Dau.; Far East: Kamch. 
Gen. distr.: Arc. Scand., Atl. Eur. Described from Spitsbergen and 
Lapland. Type in London. 

15. S. tenuis (Wahlb.) H. Sm. in Lindman, Svensk Fanerog. (l 91 8) 30.- 
S. nivalis var. tenuis Wahlb. in Fl. Lap. (1812) 1 14; Engl, et Irmsch., 
1. c, 32.- S. nivalis var.tenuior Wahlb., Fl. suec. I (l 824) 26. - 

S. stricta Hornem Oecon. PL ed. 3 (1821)470. 

Perennial; stem slender, reddish, minutely glandular in upper part, 
2.5—5 cm high; leaves in a compact radical rosette, obovate, cuneately 
tapering to petiole, crenulate, green above, reddish below, firm and rigid, 
glabrous, 0.5—0.8 cm long, 0.3—0.4 cm. broad; flowers in a capitate 
inflorescence, with very short light green pedicels or sessile, with reddish 
lanceolate bracts; calyx reddish, deeply dissected, tapering downward, with 
oblong, acuminate sepals 1— 1.5 mm long; petals white or pinkish, oval, 
with short claw, 1.5 mm long; stamens shorter than petals, with small 
anthers; pistils with long divergent styles. June — July. 

Tundra, pebbles. — Arctic : Arc. Eur. (Bol'shezemel'skaya Tundra), Nov. Z. 
Arc. Sib. (Taimyr, Yenisei mouth). Gen. distr.: Scand. Described from 
Scandinavia. 

Note. Distinguished from the closely related species S. nivalis L. 
by its size, leaf shape, and reddish color of the entire plant. 



121 



Cycle 5. Melanocentrae Engl, et Irmsch., 1. c, 46. — Inflorescence 
many -flowered, corymbiform -paniculate; petiole longer than the blade. 

16. S.tilingiana Rgl. et Til. in Nouv. Mem. Soc. Nat. Mosc. XI (1859) 
94; Engl, et Irmsch., 1. c, 57. 

Perennial; stem solitary, 10—22 cm high, glabrous, erect, leafless; 
radical leaves few (3—6), inconspicuously nerved, glabrous, the blade broadly 
ovate or rhomboid -ovate, 1.5—2 cm long, 1.2 — 1.6 cm broad, uneven-margined 
in upper part, cuneately tapering to narrow reddish petiole of the same 
length; inflorescence paniculate, ovoid -rhomboid, with oblong bracts at 
base; peduncles sparsely glandular, with small narrow bracts at their base; 
pedicels filiform, 2—4 times as long as flowers; sepals ovate or oblong, 
obtuse, divergent at anthesis;. petals white, oblong or elliptic, 3 -nerved, 
2.8—4 mm long, about twice as long as corolla, with short claw; stamens 
as long as petals; style to 2 mm long, divergent; capsule ovoid. July. 

Riverbanks. — Far East : Okh. (Ayan), Sakh. Endemic. Described from 
the Ayan River. Type in Leningrad. 

17. S. melaleuca Fisch., Cat.Hort. Gorenk. ed. I (1808) 99; Ldb., Fl. Ross. 
11,1,212; Engl, et Irmsch., 1. c, 54; Kryl., Fl. Zap. Sib. VI, 1422. - 

S. elongata Sternb., Rev. Sax. (1810) 9.— Micranthes melaleuca 
A. Los., Bull. Jard. Bot. Pr. (1928) 601.- Ic. : Engl, et Irmsch., 1. c, f. 9. 

Perennial, glabrous; stem erect, leafless, dark violet, 3—23 cm high; 
leaves petiolate, rather firm and fleshy, the blade oval-rhomboid or 
rounded -rhomboid, 1.2— 2.3 cm long, 0.8— 1 .8 cm broad, with rounded base, 
crenate in upper part, about as long as petiole; inflorescence corymbiform- 
paniculate, with small oblong bracts in the forks; calyx, pedicels, pistils, 
and stamens black-violet; calyx dissected to the middle, with triangular- 
ovate sepals; petals white or violet toward base, horizontally spreading, 
elliptic, 3 -nerved, 4—4.5 mm long, 2—2.5 mm broad, 1 .5—2 times as long 
as calyx; pistils with half-inferior flat ovary and short style; capsule 
ovoid. June— July. 

Alpine zone, rocks, stony slopes, riverbanks, tundra, near glaciers. — 
W.Siberia: Alt.; E.Siberia: Ang.-Say. Gen. distr. : N. Mong. Described 
from Sinyaya Sopka in Altai. Type in Leningrad. 



Cycle 6. Elatae A. Los. — Grex N i v a 1 i a - v i r g i ni e n s i s Engl, et 
Irmsch., 1. c, 22, p. p. — Plants with robust, strong, pubescent, many-flowered 
inflorescences and large, oblong, sinuate -dentate leaves. 

1 8. S. sachalinensis F. Schmidt, in Mem. Acad. Sc. Petersb. Se'r. XII, 7 
(1868) 133; Engl, et Irmsch., 1. c.,44. 

Perennial; rhizome with dense fibrous roots; stem erect, 15—35 cm high, 
solitary, pubescent in lower part with glandular hairs, branching in the 
inflorescence above the middle; leaves slightly fleshy, oblong-oval or oval, 
often cuneately tapering to 2 mm-broad petiole, serrate -dentate, 2—3.5 cm 
long, 1—2 cm broad, covered on both sides with rather dense minute, soft 
hairs, red -violet below, green above; panicle 5—15 cm long, oblong-ovoid 
or broadly ovoid, with scattered peduncles 1—2 cm long and filiform pedicels 



122 



rnnmiii— iillMn 



6 mm long; bracts small, oval or linear; calyx with oblong declinate sepals 
3 mm long, 3 -nerved; petals white, oval, obtuse, slightly longer than calyx; 
stamens as long as petals, with clavate filaments, capsule 5 mm long. 
July. 

Stony soils, rock crevices, seashores.— Far East: Sakh. Endemic. 
Described from Sakhalin. Type in Leningrad. 

19. S. hieracifolia Waldst. et Kit., PI. rar. Hung. I (1802) 17; Ldb., 
Fl. Ross. II, 214; Engl, et Irmsch.; 1. c.,24; Kom., Fl. Kamch. II (1929) 213; 
Kryl., Fl. Zap. Sib. VI, 1421. — Mic r a nt he s hieracifolia Haw., Enum. 
Sax. (1821) 45.- Hermesia spicata Hoppe, Taschenb. (l 800) 209. - 
Ic: Waldst. et Kit., 1. c, tab. 18. 

Perennial; stem solitary, erect, strong, thickish, 8—50 cm high, covered 
with dense, crisp, glandular hairs, leafless or with 1 small oblong leaf; 
radical leaves thickish, fleshy, oblong or elliptical-rhomboid, tapering to 
broad, short petiole or sessile, sparsely crenate,4 —8 cm long, 1—4 cm 
broad, glabrous above, pubescent below, ciliate -margined; flowers in an 
interrupted terminal, racemose inflorescence, short -branching in lower 
part, 5—10 cm long, with lanceolate, often 3 -dentate bracts longer than 
pedicels; calyx dissected to the middle into obtuse triangular sepals, 
divergent after anthesis; petals reddish or green, oval or ovate -lanceolate, 
to 3 mm long, 1 mm broad, about as long as calyx. July. 

Alpine and Arctic zones, moist moss -and -lichen tundras, marshy banks 
of mountain streams and lakes, alpine meadows near glaciers, rocks, 
solitary or in groups.— Arctic: Arc. Eur., Arc. Sib., Chuk. ; European 
part: Urals (N.); W.Siberia: Ob (N.), Alt.; E.Siberia: Ang. -Say. 
Gen. distr. : Arc. (Eur. and Am.), Centr. Eur. (mountains), Mong. Described 
from Hungary. Type in Vienna. 



Cycle 7. Stellares Engl, et Irmsch., 1. c, 66. — Stem with branching 
paniculate inflorescence; leaves oblong, cuneate, serrate -dentate; sepals 
recurved. 

20. S.foliolosa R. Br. Chlor. Melv. in Parry Journ. Voy. (l 821 ) 275; 
Turcz., Cat. Baic. (1828), No. 496; Hook., Fl. bor. Am. I (1833) 250.- 
S. comosa Fellm. in Bull. Soc. Nat. Fr. X (1863) 500.- S. stellar is $ 
comosa Retzius, Fl. Scand. Prodr. (1779); Ldb., Fl. Ross. II, 21 1 ; 
Kryl., Fl. Zap. Sib. VI, 1421; Engl, et Irmsch., I.e., 81.- S. stel laris 
v.foliosa (R. Br.) Trautv., PL imag. et descr. fl. ross. (1846) 80. — Ic: 
L., Fl. lapp. (1737) tab. 2, f. 3; Trautv., 1. c, tab. 35. 

Perennial; stem usually solitary, erect, 5—20 cm high, branching in upper 
part, with short lateral ascending branches, subglabrous, puberulent only 
in upper part, slightly sulcate; rosette leaves oblong-ovate, cuneately 
tapering, orbicular in upper part; serrate -dentate, glabrous, sparsely ciliate- 
margined, 1—2 cm long, 0.3—0.8 cm broad; inflorescence oblong or oval, 
paniculate; flowers only at tips of stem and branches, on lateral branches 
replaced by small rosettes of rather fleshy leaflets, deciduous and renewed, 
the lateral branchlets with oblong bracts at base; bracteoles at base of 
rosettes small and narrow; sepals oval, obtuse, recurved; petals white, 
lanceolate, abruptly narrowed to long claw, slightly tapering upward, 



123 



obtuse, 3 -nerved, 4— 5 mm long, more than twice as long as calyx; stamens 
shorter than petals, with dark anthers; capsule dehiscing to %, with 
short reflexed style, rounded -ovoid. August. (Plate VIII, Figure 1 1 ). 

Alpine and polar zones, river and stream banks, moist, clayey slopes, 
rocks, near snow.— Arctic: Arc. Eur., Nov. Z., Arc. Sib., Chuk., An. ; 
European part: Kar.-Lap.; E.Siberia: Yenis. (h . part), Ang. -Say., 
Dau., Lena-Kol.; Far East: Kamch., Okh. Gen.distr.: N.Am. Described 
from Lapland. 

21. S. redowskii Adams in Mouv. Mem. Soc. Nat. Mosc. Ill (1834) 241.— 
Ic: Adams, 1. c, tab. XIII, f. 2. 

Perennial; stem solitary, erect, sparsely glandular -hairy, branching 
in the inflorescence, 8—20 cm high; rosette leaves oblong or lanceolate, 
cuneate, with unequal acute teeth in upper part, 2—5 cm long, 1 .5—3 cm 
broad, glabrous below, glandular -ciliate above and along the margin; 
bracts at base of peduncles, the lower dentate, the upper smaller and 
narrow, entire; inflorescence broadly triangular -corymbiform, many- 
flowered; pedicels sparsely glandular; flowers erect, ca. 1 cm in diameter; 
calyx dissected nearly to base, with recurved, oval, obtuse sepals 3— 4 mm 
long; petals white with red dots in lower part, oblong-elliptic, 3 -nerved, 
6 mm long, 1.5 mm broad; stamens ca. half as long as petals. 
August. (Plate VIII, Figure 1 1 ). 

Forest edges, forest -tundra, tundra. — Arctic: Arc. Sib.; E.Siberia: 
Lena-Kol. (N. part). Endemic. Described from Cape Bykovskii. 

Note. Distinguished from S. s t e 1 1 a r i s L. by larger size, lesser 
pubescence, obtuse sepals, and inflorescence with greater numers of 
flowers. 

22. S. stellaris L., Sp.pl. ed. I (1753)400; Ldb., Fl. Ross. II, 21 1 

(excl. var. leucanthemifolia Misch.); Kryl., Fl. Zap. Sib. VI (l 931 ) 1421 
S. st el lar i s var. ty pi c a Engl, et Irmsch., 1. c, 71, pro max. p. — 
Spathularia stellaris Haw., Enum. Saxifr. (182 1 ) 49. 

Perennial; stems 1—5, erect, 3—15 cm high, not branching, sparsely 
pubescent; leaves stellately arranged in rosette, oblong-ovate -lanceolate, 
1— 2.5 cm long, 4— 8 mm broad, cuneate, tapering upward and mucronate, 
acutely dentate in upper part, ciliate-margined, sparsely pubescent on 
both sides; inflorescence corymbiform, few- flowered, bracts at base of 
peduncles and pedicels small, acute; pedicels minutely glandular; calyx 
dissected to base, with oblong, ovate, bluntly acuminate sepals; petals 
white, oblong-elliptic, acuminate, tapering to short claw, 3 -nerved, 5.5 mm 
long, 1.5 mm broad, 2— 2.5 times as long as calyx; stamens with short 
filaments and light anthers, shorter than petals; capsule oblong with short 
styles, dehiscing to the middle. July. 

Arctic region, moist, mossy marshy tundras, stony, pebbly soils, rocks, 
along streams.— Arctic: Arc. Eur., Nov. Z.; European part: Kar.-Lap., 
Dv.-Pech. (N.part). Gen.distr.: Arc. Scand., Greenland. Described from 
Spitsbergen. Type in London. 



Cycle8. Merckianae Engl, et Irmsch., 1. c, 88. — Loosely cespitose 
plants; stems few-flowered; pistils with subsessile stigmas. 



124 



m^^^^^M 



160 



23. S. merckii Fisch. in Sternb., Rev. Suppl. (1822) 1; Ldb., Fl. Ross. II, 
208; Engl, et Irmsch., 1. c, 88; Kom., Fl. Kamch. Il(l92 9)215.-Leptasea 
merckii (Fisch.) Kom., 1. c. — Ic. : Sternb., 1. c, tab. 1 . 

Perennial, forming loose flat tufts; rhizome with slender woody shoots; 
roots slender, fibrous; stems erect or ascending, densely glandular, 
4—10 cm high; rosette leaves often remote, oboval or lanceolate, usually 
acuminate, less often orbicular, tapering downward, glabrous below, sparsely 
hispid above, densely hispid on the margin, sometimes obscurely 3 -dentate 
in upper part, 1—2.5 cm long, 0.4—0.8 cm broad; flowers 1—5, 1—1 .5 cm in 
diameter, their long pedicels with small lanceolate bracts at base; calyx 
dissected to the middle, glabrous, with oval, obtuse, ciliate-margined sepals; 
petals white, rounded, 5 -nerved, long-clawed, 0.4—0.5 cm long, more than 
twice as long as sepals; stamens longer than calyx, shorter than petals, 
with slender yellowish filaments and small dark purple anthers; stigmas 
subsessile. July. 

Loose soils, sands, products of volcanic eruptions, in great masses 
along mountain stream banks in the alpine region.— Arctic: Arc. Sib., 
An.; Far East: Okh., Kamch. Endemic. Described from the Yablonovy 
Range. Type in Leningrad. 



Section 2. HIRCULUS (Haw.) Tausch, Hort. Canal, (l 823) 1.- Hir cuius 
Haw., Enum. Sax. (1821 ) 40 (pro genere). — Leptasea Haw., Enum. Sax. 
(1821) 39 (pro genere); Small, N. Amer. Fl. XXII, 2 (1905) 151 (excl. non- 
nullis specibus). — Ovary superior, not imbedded or else hypanthium well- 
developed; capsule with divergent syle, dehiscing to Y 3 — x / 2 ; petals yellow; 
cauline shoots reduced; flowering stems leafy; leaves entire, pubescent, 
hispid, or glandular, or with cartilaginous margin. 



Cycle 1. Hirculoideae Engl, et Irmsch., 1. c, 106. — Stems pubescent, 
eglandulose; leaves glabrous. Not forming rooting shoots. 

24. S.parnassioides Rgl. et Schmalh., Descr. PI. nov. rar. O. Fedtsch. 
in Turk. lect. in Izv.O-va Lyub. Ect. Ill, 1 8 (1882)27; Engl, et Irmsch., 
I.e., 126. 

Perennial; stem slender, 8—12 cm high, covered with scattered articulate 
hairs; radical leaves unknown; cauline leaves 2, small, 1.5 cm long, 1 cm 
broad, oval, mucronate, entire, glabrous, 3 -veined, short -petioled; flowers 2, 
with pedicels to 2.5 cm long, the upper flower terminal, the other in axil 
of first leaf; sepals oblong, obtuse, 3—4 times as long as fused part of calyx; 
petals oboval, 3 -nerved, white, more than twice as long as calyx; styles 
erect, with capitate stigmas. July. 

Centr. Asia: Pam.-Al., collected in the Isfairam River valley. Endemic. 
Type unknown. 

Note. Apparently no herbarium specimen has been preserved. The 
initial description was made after a single specimen from the collections 
of O. A. Fedchenko, from which a drawing was made; this is in the 
herbarium of the Biological Institute. 



125 



25. S. hirculus L., Sp.pl. ed. 1 (1753)402; Ldb., Fl. Ross. II, 210; 
Ldb., Fl. Alt. II, 121; Engl, et Irmsch., 1. c, 1 10 ; Kryl., Fl. Zap. Sib., VI, 
1425; Kom., Fl. Kamch. II (1929) 207. - S. nutans Adams in Nouv. Mem. 
Soc. Nat. Mosc. Ill (1834) 242 (?).- Hirculus ranunculoides Haw., 
Enum. Sax. (1821 ) 40. - L e p t a s e a hirculus (L.) Small, N.Am. Fl. XXII 
(1905) 152.- Exs.: HFR, No. 19. 

Perennial; stems simple, erect, solitary or several and then plants 
cespitose, leafy, with a basal leaf rosette, sometimes with leafy, nonrooting 
shoots, subglabrous in lower part, more or less densely covered with 
rufous hairs in upper part below inflorescence; leaves light green, entire, 
lanceolate, 1 — 3 cm long, 3—5 cm broad, glabrous, the lower tapering to 
pubescent, rather long petiole, the upper sessile, narrower and smaller; 
flowers 1—4, at tip of stem; calyx dissected nearly to base, the sepals 
oblong, apically rounded, ciliate -margined, reflexed after anthesis, 
2.5 — 5 mm long; petals bright yellow, sometimes with orange dots, elliptic, 
8—12 mm long, 3—3.5 mm broad, with convex nectary below; stamens as long 
as petals; pistil divided only at apex into styles; capsule oblong -ovoid, 
1 cm long. August — September. 

Forest, Arctic, and alpine regions, moist moss -and -lichen tundras, along 
small rivers and streams, moist meadows. Hyp nu m and Sphagnum 
bogs. — Arctic : Arc. Eur., Nov. Z., Arc. Sib., Chuk., An.; European part : 
Kar.-Lap., Lad.-Ilm., Dv.-Pech., V.-Don., U. Dnp., U. V., V. -Kama, M. Dnp., 
L. Don (N. part); Caucasus: Dag., E. and S. Transc; W.Siberia: Yenis., 
Ang.-Say., Lena-KoL, Dau.; Far East : Ze. -Bu., Okh., Kamch. ; Centr.Asia: 
Pam. -AL, T. Sh. Gen. distr. : Arc. Eur., Centr. Eur., Scand., Atl. Eur. 
Described from Switzerland. Type in London. 

Note. Size, height of stem, size and number of flowers vary greatly; 
two forms can be differentiated: f . m a j o r Engl, et Irmsch, with large 
stems, 1—4 flowers, not cespitose, a form confined to the forest zone and 
the lower mountain zone; and f. minor Engl, et Irmsch., a cespitose 
1 -flowered form, with large flowers, stems often numerous, 4—10 cm high, 
confined to the Polar-Arctic region, and the high-mountain zone. There 
are transitions between the two forms. 



Cycle 2. Flagellares Engl, et Irmsch., 1. c, 1 57. — Leaves glandular 
or ciliate; stems with above-ground rooting shoots. 

26. S. flagellaris Willd. ex Sternb., Rev. Sax. (l 810) 25; Ldb., Fl. Ross. II, 
209 p. p.; Kom., FL Kamch. II (l 92 9) 2 16. - S. a s p e r a M. B. Fl. taur. -cauc. I 
(1808) 314.- S. sobolifera Adams in Nouv. Mem. Soc. Nat. Mosc. Ill 
(1834) 243. — S. f la ge 11a r i s var. s t e n o s e p a la Engl, et Irmsch., 1. c, 
159.- Exs.: Fl. cauc. exs., No. 135. 

Perennial; stem solitary, erect, simple, with evenly spaced leaves, 
glandular -pubescent, 5—20 cm high, giving rise at base to slender, above- 
ground, glabrous, creeping shoots, 3—12 cm long, terminating in a subsequently 
rooting bud; leaves entire, ciliate -glandular on the margin, glabrous above, 
sometimes glandular -pubescent below, radical leaves rosulate, oblong- 
obovate, tapering to petiole; cauline leaves elliptic, slightly tapering toward 
base, sessile, all leaves acute, to 1 5 mm long and 5 mm broad; flowers 



126 



wMm—ma^^n^^^—^—^^^mmam^mm 



1-5 at tip of stem, with pedicels of equal length (1-2 cm long), broadly 
campanulate, 15 — 18 mm in diameter; calyx dissected to base, glandular, 
the sepals narrowly elliptic, acute, 2— 4 mm long, 1 — 1.5 mm broad; 
petals yellow, broadly oval-ovate, 8—13 mm long, 4—6 mm broad; stamens 
half as long as petals; capsule rounded -ovoid. .June — August. (Plate IX, 
Figure 11 ). 

Alpine and Arctic regions, taluses, sands, pebbly tundra, stony and pebbly 
soils, along rivers, gorges. — Arctic : Arc. Sib.; E.Siberia: Dau., 
Ang -Say.; Far East: Kamch., Caucasus: Cisc. (Main Range), Dag.; 
Gen. distr.: Ber. (Alaska), Mong. (Kentei, Kosogol). Described from the 
Caucasus. Type in Berlin. 

2 7. S. komarovi A. Los. sp. nova in Addenda VIII, p. 372.— S.flagellaris 
auct. Fl. As. Med.; Fedchenko, Rast. Turk. (1915) 480.- Exs. : HFR, No. 69 
(sub S.flagellaris Willd.). 

Perennial, loosely cespitose; stems covered by dead leaves, 4—7 cm 
high, strongly pigmented, rather sparsely glandular, leafy, with slender 
above-ground shoots bearing rooting buds; leaves fleshy, reddish, slightly 
concave, oval-lanceolate, bluntly acuminate, slightly tapering downward; 
radical leaves in a dense rosette, to 1 cm long, 4 mm broad; cauline leaves 
diminishing toward tip of stem, narrower; bracts pedicellate, sublinear, 
all leaves glandular on the margin and on both sides; flowers (l— 2)3— 8 
(mostly 6) in a corymb, erect, campanulate, 8— 9 mm in diameter, with 
glandular pedicels about half as long as stem; petals obovate, yellowish 
pink, tapering downward, rounded at the apex, 6 — 8 mm long, 3—4 mm broad, 
3-nerved; calyx dissected to base, broadly cylindrical, with rounded -truncate 
base, the sepals erect, oblong, obtuse, pigmented, 2—4 mm long, 1 — 1 .5 mm 
broad, concave in lower part, separated by an obtuse notch; pistils with 
oblong ovary and subsessile stigmas. July— August. (Plate IX, Figure 10). 

Passes, slopes, glacial moraines, on clayey and pebbly soils.— Centr.Asia: 
Pam.-Al. (Zeravshan, Shugnan). Endemic. Described from the Zeravshan 
Range. Type in Leningrad. 

Note. Note should be taken of the smaller-flowered and less pigmented 
Shugnan form. 

28. S. setigera Pursh, Fl. Am. Sept. I (1814) 312.- S.flagellaris 
var.setigera Engl., Mon. Sax. (l 872 ) 225. - S. f la g e 1 la r i s var. 
platysepala Engl, et Irmsch., 1. c, 15 9. — S. myosotidiflora Don in 
Transact. Lin. Soc. XIII (1821 ) 373.- Pall, ex Spreng.,Syst. Veg. II (1825) 
364.- S.flagellaris Ldb., Fl. Ross. IV, 209, p. p.; Kryl., Fl. Zap. Sib. VI 
(1931) 1427.- Ic: Ldb., Ic.pl. Fl. ross. tab. 321; Holm., Nov. Sem. Veg. 
(1885) tab. 9, f. 1-7. 

Perennial; stem solitary, erect, 2—15 cm high, strongly glandular, 
pubescent, with evenly spaced leaves, giving rise at base to above-ground 
slender, glabrous shoots terminating in rooting buds; leaves entire, short- 
acuminate, ciliate, densely glandular on both sides, 5 — 15 mm long, 2.5 mm 
broad; cauline leaves sessile, lanceolate; radical leaves rosulate, oblong- 
ovate, tapering to broad petiole; flowers 1—5 on very short, densely glandular 
pedicels, campanulate -infundibular, with distinctly developed hypanthium, 
10 — 15 mm in diameter; calyx dissected to the middle, 6—8 mm long, densely 



127 



glandular, with oval, obtuse sepals; petals yellow, obovate, 8—9 mm long, 
5—6 mm broad; pistils with half-inferior ovary, dissected to % into 
styles; capsule oblong. July — August. (Plate IX, Figure 12). 

Alpine and Arctic zones, stony moss -and -lichen tundra, alpine meadows. 
Arctic: Nov. Z., Arc. Sib.; W.Siberia: Alt.; Centr.Asia: T. Sh. Gen. distr. 
Arc. Eur., N. Am. Described from Cape Newenham, Arc. Am. 



Cycle 3. Sedi formes Engl, et Irmsch., 1. c. — Plants with creeping, 
leafy cauline shoots, cespitose. 

29. S. serpyllifolia Pursh, El. Am. Sept. I (1814) 311 ; Ldb.. El. Ross. II 
(1844) 210; Engl, et Irmsch., I.e., 151; Kom., Fl. Kamch. II (1929) 216.- 
S. fischeri Ser. in UC.,Prodr.IV (1830) 22.- S. b i c o 1 o r Stcrnb., 
Suppl. II (1831)49, tab. 14.— Leptasea serpyllifolia (Pursh) Small, 
N.Am. Fl. XXII (1905) 152.- Ic: Engl., Rot. Jahrb. XLV1II (1912) f. 15. 

Perennial; loosely cespitose; cauline shoots creeping, leafy, woody; 
flowering stems 2—5 cm high, oligophyllous, erect, glandular; leaves 
spatulate, tapering downward, obtuse, with thin cartilaginous margin, dark 
green, glabrous, 6—7 mm long, 1 .5—2 mm broad; cauline leaves smaller, 
often emarginate; flowers 1—2, erect; calyx incised to 2 / 3 into broadly oval- 
obtuse, glandular sepals, reflexed toward end of anthesis; petals golden- 
yellow, broadly oboval, with short claw, 5—6 mm long, 3—4 mm broad, 3 times 
as long as calyx; stamens much shorter than petals; capsule ovoid, with 
divergent styles. July. 

Rock streams, dry alpine meadows, rocks. — Arctic: Arc. Sib., Chuk., 
An.; E.Siberia: Yenis., Lena-Kol. (N. part); Far East: Kamch. Gen. distr. : 
Ber., N.Am. Described from N.America. 

Note. Ledebour reports this species for the Altai, according to the 
collections of Pallas, which are also mentioned by Engler, but there is 
no material from the Altai in the Leningrad herbarium; therefore, 
Ledebour's indication should be considered erroneous. 



Section 3. MISCOPETALUM Haw., Syn. PL succ. (1812) 323; Ej., Enum. 
Sax. (1821) 16.- Micropetalum Tausch, Hort. Canal. I (1823) 1.- 
Ovary free, not sunken; calyx with flat base; capsule oblong-ovoid, with 
short styles. Stems leafy; radical leaves long-petioled, the cauline 
short -petiolcd, all leaves dentate. 

30. S. coriifolia (Somm. et Lev.) Grossh., Fl. cauc. II (1932)235.- 
S. rotundifolia var. coriifolia Somm. et Lev., Enum. in A. H. P. XVI 
(1900) 175; Oetting., in Fl.cauc. crit. 40 (l 91 3) 20.- S. rotundifolia Ldb., 
Fl. Ross. II (1844) 217, non L. - Exs. : Broth., PL cauc, No. 372; Sintenis 
It. or., No. 3697. 

Perennial; stem erect, to 50 cm high, sulcate, glandular -hairy; branching 
in inflorescence; radical leaves rosulate, rather numerous, with pubescent, 
rufous petioles 2—3 times as long as the blade; blade rounded -reniform, to 
6 cm long, 5 cm broad, coriaceous, with unequal, broad, acuminate or bluntly 
acuminate teeth, sparsely hairy on both sides or glabrous, ciliate-margined. 



128 



165 



often red -purple below; cauline leaves small, with narrow, acute teeth; 
inflorescence sparse, spreading, ova] in outline; flowers 3 or 4 at ends 
of branches, their pedicels with varying Length, with small, narrow 
bracteoles; inflorescence branches and pedicels densely glandular; flowers 
12 — 15 mm in diameter; calyx dissected to base, glandular, with oval-oblong 
subobtuse sepals; petals white with red dots, less often pure white, oblong, 
narrow, tapering to small claw, with rounded tip, 3 -nerved, 7-8mm long, 
2—2.5 mm broad, twice as long as calyx; stamens shorter than petals, 
as long as pistils; pistils with short, divergent styles; seeds Light brown. 
May — June. 

Moist soils in the shade in forests to the subalpine zone.— Caucasus: 
W. and E. Transc. Gen.distr.: [ran.. Arm. -Kuril. Described from the 
Batumi District, between Batumi ami Ad/hari -Tskhali. Type in Florence. 



Section 4. CYMBALARIA Griseb., Spicil. I'M. Rumel. I (l 843) 336. - 
Ovary free, not sunken; calyx with flat base; capsule with short styles; 
seeds round. Annual plants with leafy, branching stems and thin spatulate 

leaves. 

31. S. huetiana Boiss., Diagn. PI. or. Sen 2, fasc. 11 (1856) 72; I'M. Or. 
(1872) 812; Oetting. in Fl. cauc. crit. Fasc. 42 (1915) 11.- S. cy mbala r ia 
var. huetiana Engl, et Irmsch., 1. c. 

Annual; stem glabrous or sparsely glandular-pubescent, ascending, 
nodding or decumbent, branching; leaves thin, light green, with very 
conspicuous light brown glands below; lower leaves with petioles twice as 
long as the blade, cordate -reniform or with truncate base, to 2.5 cm 
broad, 2 cm long, 5— 7-lobed, the lobes broadly oval, rounded or subacuminate 
upper leaves smaller, 3— 5-lobed, often with cuneate base, short -petioled; 
pedicels slender, long, erect; sepals triangular -oval, acute, reflexed; petals 
as in the preceding species. May— July. (Plate IX, Figure 1 ). 

Subalpine meadows, stream banks, moist rocks. — Caucasus: W. and 
E. Transc, Cisc. Gen.distr.: Arm. -Kurd. Described from the vicinity 
of Trebizond. Type in Geneva. 

N ot e. Little studied species; very little is known concerning habitat. 

32. S. cymbalaria L., Sp. pi. ed. I (1753)405; Ldb., Fl. Ross. II, 223; 
Boiss., Fl. Or. II (1872) 811.- S. orient a lis Jacq., Obs. bot. 11 (1767) 9, 
tab. 34.- S. par ad ox a M. B. Fl., taur.-cauc. I (1808) 317; 111 (l 81 9) 2 95, 
non L. — Lobaria ori en talis Haw., Enum. Sax. (1821 ) 18. — 

S. cymbalaria var. e u c y m b a 1 a r i a Engl, et Irmsch., 1. c, 202. — 
Exs.: HFR, No. 361; Fl. cauc. exs., No. 264. 

Annual; stem ascending, erect or prostrate, to 30 cm long, profusely 
branching, leafy, glabrous or sparsely pubescent; leaves flat, somewhat 
fleshy, light green, glabrous or glandular -hairy, the lower leaves with 
petioles twice as long as the blade, the upper shorter-pctioled, all leaves 
reniform or rounded -reniform, the upper sometimes broadly oval with 
cuneate base, 5—25 mm long, 8—35 mm broad, with 5—13 large tringular-oval 
acute teeth, the middle tooth larger than the others, more obtuse, and often 
slightly cordate at base; flowers in axils of upper leaves and at ends of 



129 



166 



branches, with long, slender, glandular pedicels reflexed after anthesis; 
calyx dissected from base, with oval -triangular subobtuse sepals, 
1 .5—2 mm long, 0.5—0.8 mm broad, horizontally spreading or divergent, 
glabrous or glandular; petals oblong-elliptic, to 6 mm long, 2 mm broad, 
3 -nerved, subobtuse, with short claw, white or pinkish-yellowish with 
orange spot; stamen half as long as petals; ovary as long as sepals; 
capsule ovoid, to 4 mm long; seeds round. May. (Plate IX, Figure 2). 

Forests of the upper forest zone and at the timberline, rocks in gorges, 
on limestones, subalpine meadows. Caucasus: W. and E. Transc, Lenkoran. 
Gen. distr. : Arm. -Kurd., Iran. Described from the East. Type in London. 



Section 5. TRIDACTYLITES Haw., Enum. Sax. (1821 ) 21 . - Flowers 
small; ovary semi-imbedded; calyx dissected to 1 / 3 — 1 / 2 , with erect sepals; 
capsule with short styles. Annuals or biennials with erect leafy stems and 
radical rosette of 3—5 quinquelobate leaves. 

33. S.tridactylites L., Sp.pl. ed. I (1753) 404; Ldb., Fl. Ross. II, 226; 
Shmal'g., Fl. Sr. i Yuzhn. Ross. I (1895) 355.- S.trifida Gilib., Fl. Lith. 
(1781) 181.- S. annua Lap., Fl. Pyr. (l 801 ) 597. - S. t r id a c t y li t e s var. 
minor A. Blytt, Norg. Fl. Ill (1876) 910.- S.tridactylites subsp. 
minor A. Blytt, Handb. i Norg. Fl. (l 906) 410. - S. t r i d a c ty 1 it e s subsp. 

e ut r id a c ty 1 it e s Engl, et Irmsch., Pflanzr., 1. c, 206. — T r i d a c t y 1 i t e s 
annua Haw., Enum. Sax. (l 821 ) 21 . - Exs. : HFR, No. 20. 

Annual, completely covered with small glandular hairs; stem 2 — 18 cm 
high, erect or flexuous, usually slender, less often thickish, simple or 
branching from the very base, sparsely leafy; radical leaves rosulate, 
sometimes absent, oblong -spatulate, cuneate, often trilobate, to 2 cm long, 
0.5 cm broad; cauline leaves smaller and narrower, 3— 5-lobed, sometimes 
entire (in low-growing forms); inflorescence spreading, with slender 
pedicels, twice as long as flowers or longer, elongating after anthesis; 
first flowers bisexual, the following ones with obsolete pistils and stamens; 
calyx dissected to the middle, with subobtuse or orbicular sepals, to 2 mm 
long; petals oval-cuneate, white, more than twice as long as calyx; ovary 
half-sunken, hemispherical, with rounded base ; stamens as long as sepals, 
attached to their base; pistils with short widely-scattered styles; capsule 
globose. May — June. (Plate IX, Figure 4). 

Shores, limestones, rocks. — European part : Lad.-Ilm., U. Dnp., Bl., Crim. : 
Caucasus: Cisc, Dag., W., S., and E. Transc. ; Centr. Asia: Mtn. Turkm. 
Gen. distr.: Centr. Eur. ,Scand.,W. Med. , Bal.-As. Min. Described from Europe. 

Note. Species varying greatly in overall size, leaf incision and flower 
dimensions. Litvinov has noted for Turkmenia f. simplex Litw., with 
very small flowers and slender stem, scarcely branching. Engler reports 
9 forms differing in their growth in the same distribution area. 

34. S.adscendens L., Sp. pi. ed. I (1753) 405. — S. c ont r over sa Sternb., 
Rev. Sax. (1810) 43. — S. t r idacty lit es subsp. ad s c e nd e n s Engl, et 
Irmsch., 1. c, 217.- Exs.: FL exs. Reipubl. Slov., No. 225 (1926). 

Annual, completely covered with dense glandular hairs; stem erect, to 
25 cm, usually branching in upper part; leaves in dense radical rosette; 



130 



cauline leaves evenly spaced throughout the stem, elliptic -cuneate, entire 
or trilobate; radical leaves to 2 5 mm long, 7 mm broad, the cauline narrower 
and smaller; inflorescence 1 - to many -flowered, broadly paniculate; 
pedicels slender, 3—4 times as long as flowers, curved in upper part; 
calyx campanulate, dissected to V 3 into acute triangular sepals tapering 
toward base; petals 3—4 times as long as sepals, white, to 4 mm long, 
3 mm broad, 3 -nerved, obovate; stamens and pistils as in preceding species. 
May- July. (Plate IX, Figure 3). 

Riverbanks, rocks, pebbles, on moist soil.— Caucasus: Dag., W. and 
E. Transc. Gen. distr. : Centr. Eur., Atl. Eur., Scand., W. Med., Bal. -As. Min. 
N.Am. 

Note. Varying greatly, like the preceding species. Engler reports 
several forms within the species, but — as in the preceding species — these 
forms differ mainly in their ecology. 



Section 6. NEPHROPHYLLUM Gaud., Fl. Helv. Ill (1828) 85 (excl. 
S. rotundifolia). — Sobaria Haw., Enum. Sax. (l 821 ) 20 p.p. (pro gen.). 

Flowers large, campanulate, with white petals, several times as long 
as calyx; ovary slightly imbedded; styles slender, divergent. Leaves 
large, dentate or lobate, reniform or oblong-rounded, cuneately tapering. 
Stem leafy, usually petiole bears at its base bulbil-tubercles. 



Cycle 1. Irriguae Engl, et Irmsch., 1. c, 234. — Plants with bulbils 
at base of petioles; leaves twice trilobate. 

35. S.irrigua M.B. Fl.taur.-cauc. II (1808) 460, III (1819) 295; Ldb. 
FL Ross. II, 218; Shmal'g., FL Sr. i Yuzhn. Ross. I, 355; Engl, et Irmsch., 
1. c, 2 34.- S.aquatica M. B. Fl.taur.-cauc. I (1808) 317, excl. syn. Lap. - 
S. petraea Hablizl, Phys. Beschr. d. Taur. Statt. (1789) 257, non L. - 
S. r a nunc uloid e s Haw. Enum. Sax. (1821 ) 25. — S. g e r a n i od e s /3 
irrigua Ser. in DC. Prodr. IV (1930) 30. - Ic. : Bot. Mag. tab. 220. 

Perennial, completely densely soft -hairy; stems numerous, 8—85 cm 
high, sulcate, often pinkish, thick, branching in upper part; radical leaves 
numerous, long-petioled, rounded -reniform in outline, trilobate, each lobe 
dissected into oblong, subobtuse, rarely acuminate lobules; blade 0.5—5 cm 
in diameter, on the average mostly 2 cm; petioles pinkish, sulcate, thickish, 
2—4 times as long as the blade; cauline leaves short -petioled or sessile, 
smaller, with narrower acute lobules; bracts to 1 cm long, lanceolate; 
flowers erect, in dense, corymbiform-paniculate inflorescence, with rather 
slender, short pedicels; calyx with narrowed base, dissected nearly to base 
into linear obtuse sepals to 5 mm long, 1 mm broad, 3 -nerved; petals white, 
oblong -ovate -cuneate, 10 — 12 mm long, 3— 4mm broad, 3 -nerved; stamens half 
as long as petals; ovary semi -imbedded, with filiform erect styles as long as 
stamens; capsule ovoid. May. (Plate K, Figure 7). 

Mountain slopes, rocks, bluffs. — European part: Crim. Endemic. 
Described from the Crimea. Type in Leningrad. 

Cycle 2. Granulatae Engl, et Irmsch., 1. c, 236. — Plants with bulbils 
at base of petioles. Ovary sunken. 



131 



36. S.granulata L., Sp.pl. ed. 1 (1753)403; Ldb., Fl. Ross. II, 21 8; 
Shmal'g., Fl. Sr. i Yuzhn. Ross. I (1895) 355; Engl, et Irmsch., 1. c, 245 
(sub eugranulata). — S.corymbosa Luce, Prodr. Fl. osil. (l 823) 1 37. — 
S. carnosa Luce, 1. c, 137.- Ic: Rchb., Ic. Fl. Germ, et Helv. XXIII, tab. 
tab. 116. 

Perennial; stem erect, solitary, leafy in lower half, scabrous -hairy, 
15— 50 cm high, branching in upper half; radical leaves in dense rosette, the 
long petioles twice as long as blade, covered with white flexuous hairs, 
the blade rounded -reniform, 1—5 cm broad, 0.7—2.5 cm long, 5 — 9-lobed, 
the lobes rounded, glandular -hairy on both sides; cauline leaves with 
cuneate base and narrower, more acute lobes, short -petioled; bracts 
deeply dissected into 3—5 acute lobes, or entire, lanceolate; flowers in 
many -flowered, broad, corymbiform -paniculate inflorescence, large, erect, 
with glandular pedicels of varying length, sometimes subsessile; calyx 
deeply dissected into lanceolate, acute sepals, glandular, to 5 mm long; 
petals white or yellowish, oblong-spatulate, broadening in upper part, 
cuneately tapering downward, 10—14 mm long, 3—5 mm broad; stamens 
half as long as petals; ovary 3 / 4 imbedded, with styles as long as stamens; 
capsule ovoid, to 7 mm long; seeds black, oblong, to 0.5 mm long. May. 
(Plate IX, Figure 8). 

Meadows, ditches, meadow slopes and hills in sparse forests.— 
European part : Lad. -Ilm., U. Dnp. Gen. distr. : Centr. Eur., Scand., 
W. Med., Bal. -As. Min. Described from Europe. Type in London. 



Cycle 3. Sibiricae Engl, et Irmsch., 1. c, 262. — Plants with bulbils 
at base of petioles; ovary almost free. 

37. S.cernua L., Sp.pl. ed. I (1753) 403; Ldb., Fl. Alt. II (l 830) 122 ; 
Engl, et Irmsch., I.e., 270, f. 64; Kom., Fl. Kamch. II (1919) 205.- 
S.bulbifera Gunn., Fl. Norveg. II (1772) tab. 8. - S. simulata Small 
in N. Amer. Fl. XXII, 2 (1905) 128.- Ic. : Rchb., Ic. Fl. Germ. XXII, tab. 118. 

Perennial; underground shoots covered with fleshy petioles of obsolete 
leaves, forming buds; stems solitary, 25—35 cm high, erect, branching 
or simple (f. simplicissima Ldb., Fl. Alt. II (1830) 122), slightly 
glandular -pubescent, with leaves evenly spaced and bearing small reddish 
bulbils in leaf axils; radical and cauline leaves petiolate, to 3.5 cm in 
diameter, reniform, with 5—9 large acute, broadly oval or narrowly triangular 
lobes; in upper leaves, the lobes more acute and narrower; bracts with 
bulbils in their axils, entire, oval or lanceolate; flower solitary at tip 
of stem, with rather long, glandular pedicels, large, campanulate; sepals 
oval, not acute, pubescent, 2.5— 3.2 mmlong; petals white, oblong-oval, 
3 -nerved, tapering downward, without claw; stamens as long as calyx; 
ovary half -inferior, with short styles and rather large stigmas; throughout 
the stem are bulbils — buds with numerous, fleshy, reddish leaflet lobes, 
deciduous and renewed. May — July. 

Forest and Arctic zones, rocks, pebbles, alpine meadows. — Arctic: 
Arc. Eur., Arc. Sib., Chuk., An.; European part : Kar. -Lap., Dv. -Pech., 
V. -Kama (N.Urals); W.Siberia: Alt., Ob (N. part); E.Siberia: Yenis. 
(N. part), Dau., Lena -Kol., Ang. -Say; Far East : Kamch; Centr. Asia: T. Sh. 
Described from Lapland. Type in London. 



132 



(169) 




PLATEIX 1-Saxifraga huetiana Boiss.,leaf; 2 - S. cy mbalari a L.,leaf; 3 - S.ad see nd ens L. 
a) flower; 4 - S. ridacty li tes L. ; 5 - S. mollis Smith, a) leaf, b) petal; 6 - S.sibiri ca L., petal; 
7-S.irrigua M.B.,leaf; 8 - S. g r a n u la ta L., leaf; 9 - S. r i v ul a r i s L.; 10 - S.komarov i A. Los., 
a) leaf, b) flower; 11 - S. f lagella r is Willd.. a) leaf, b) flower; 12 - S.se t i ge ra Pursch., 



a) leaf, b) flower. 



133 



38. S. sibiricaL., Sp.pl. ed. II (1762) 577; Ldb., Fl. Ross. II, 219; Boiss., 
Fl.Or. II, 807.-S. granulata Schangin in Pall.n. nord. Beitr. VI (1793) 
93. -S. cernua ,8 . s i b i r i c a Korshinsky, Tent. Fl. Ross. Or. (1898) 164.- 
S. stephaniana Sternb., Rev. Suppl. I (1822) 8, tab. 6, f . 2 . - S. g r a n d i - 
flora Sternb., Rev. (1810) 20, tab. 12, f. 4.-Lobaria sibirica Haw., 
Enum. Sax. (1821) 21. — S. sibirica L. v.eusibirica Engl.et Irmsch., 
I.e., 263.- Ic: Sternb., Rev. (1810) tab. XII, 4 sub S. granulata. - 

Perennial; stem erect, 4 — 30 cm high, leafy, pubescent in upper part, with 
small whitish underground tubercles at base; radical leaves reniform, 
6 — 20 mm long, 8 — 25 mm broad, incised nearly to the middle into 7 — 9, 
less often 5 broadly ovate, obtuse, orbicular or acuminate lobes, glabrous 
and glandular on the margin, long-petioled, the middle cauline leaves 
shorter-petioled, smaller, trilobate, the uppermost entire or lobate; flowers 
2 — 7 at tip of stem, the slender, long pedicels with small ovate-lanceolate 
bracts; calyx glandular-pubescent, dissected to three-quarters into oblong or 
ovate sepals; petals white, 5 — 7-nerved, obovate, cuneate toward base, 2 — 3 
times as long as calyx; ovary superior. June — July. (Plate IX, Figure 6). 

Stony, pebbly slopes, rock crevices, sparse forests in the alpine and 
forest zones, less often in the steppe region.— European party. V. -Kama, 
Transv.; W.Siberia: U. Tob., Irt., Alt.; E.Siberia: Ang. -Say ., Yenis ., 
Dau., Lena-Kol.; Far East: Ze.-Bu.; Centr.Asia: T. Sh., Pam. -Al., Syr . D., 
Dzu.-Tarb. Gen. distr.: Dzu. -Kash., Mong. Described from Siberia. 
Type in Leningrad. 

Note. A form with small sessile cauline leaves, slender stems and 
1 — 3 small flowers has been observed in the Shugnan and the Pamir. 

39. S. mollis Smith in Sternb., Rev. Suppl. II (1832) 37. - S. cymbalaria 
M. B., Fl.taur. -cauc.III (1819) 202 (excl. syn. Willd.).- S. sibirica 

auct. Fl.cauc- Ic: Sternb., 1. c, tab. 25, f . 1 . - Exs.: Kotschy 1859 No. 521; 
Sintenis, Her. or. 1894 No. 5664. 

Perennial; stems solitary, with underground tubercles at base, 10— 30 cm 
high, glandular-pubescent, often lilac, erect or ascending; leaves reniform, 
with 5 — 7 large, triangular-ovate, acuminate lobes, 1 — 2.5 cm long, 1 — 3 cm 
broad, the lower with long, glandular-pubescent petioles, the middle leaves 
shorter-petioled, the upper subsessile, the uppermost broadly ovate, acute; 
all leaves with glabrous blade, glandular on the margin, dark-veined; 
flowers few, in a corymb, the slender, glandular pedicels with small 
lanceolate bracts; calyx subobtuse, dissected to base; sepals sublinear, 
bluntly acuminate, to 0.5 cm long, 1mm broad, glandular; petals white, 
oblong-obovate, tapering to long claw, 3-nerved, 2 cm long, 0.6 cm broad; 
stamens shorter than petals, with small spherical anthers; ovary with short 
style. June — July. (Plate IX, Figure 5). 

Stony soils, moraines, limestone taluses in the alpine zone.— Caucasus: 
Cisc, S., E., and W. Transc. Gen. distr.: Iran., Arm. -Kurd., As. Min. 
Described from Mt. Ararat. 

Note. Distinguished from S. s i b i r i c a L. by the subtruncate base of 
calyx, longer and correspondingly narrower petals with 3 nerves instead 
of 5, the entire plant more delicate and less leafy. 

40. S.rivularis L., Sp.pl. ed. I (1753) 404; Ldb., Fl. Ross. II, 221 ; Engl, 
et Irmsch., I.e., 277; Kom., Fl. Kamch. II (1929) 206; Kryl., Fl. Zap. Sib. 



134 



173 



VI (1931 ) 1413. — S. cymbalaria vel nov. sp. Chamisso in Linnaea VI 
(1831) 555. -Ic: Sternb., Rev. Sax. (1810) 812, f. 3. 

Perennial; stems 3—10 cm high, numerous, erect, leafy, covered with 
white hairs, with underground shoots; leaves light green, somewhat fleshy, 
palmately 3 — 5-lobed, the radical long-petioled, the cauline short-petioled, 
entire, oblong, sessile; stipules broad, ciliate; flowers in leaf axils 
throughout the stem, the middle flowers shorter, the upper sessile; calyx 
glandular, dissected to the middle, with oblong sepals; petals 3 — 4 mm long, 
white, obovate or oblong-ovate; capsule globose, to 4 mm long, with divergent 
short styles; calyx and capsules often purple. July — August. (Plate IX, 
Figure 9). 

Polar-Arctic and alpine regions, pebbles, tundra spring banks, meadows, 
rarely in the mountain forest belt, always in the shade. — Arctic: Arc. 
Eur., Nov. Z., Arc. Sib., Chuk., An.; Far East: Kamch. Gen. distr .: Arc, 
Scand., N.Am. Described from Lapland. Type in London. 

41. S.bracteata D.Don in Transact. Linn. Soc. XIII (1822) 367; Kom., 

Fl. Kamch. II (1929) 207. - S. laurentiana (Ser.) Engl . in Engl.et Irmsch., 
I.e., 282. — S. cymbalaria Cham, in Linnaea VI (1831) 555, non L. 

Perennial; stem 6 — 15 cm high, erect, usually solitary, pubescent with 
whitish or rufous hairs, subtomentose in upper part, slightly sulcate; 
leaves broadly reniform, 1—3 cm long and broad, with 3 — 7 large, rounded 
lobes, lower and radical leaves long-petioled, the upper short petioled, the 
uppermost sessile, approximate to flowers; petioles flat, rather fleshy, 
the lower sparsely pubescent, the upper rather densely hairy; flowers 1 cm 
in diameter, few, in axils of bracts, with very short pedicels, forming a 
condensed subcapitate inflorescence; calyx dissected to the middle, 
subglabrous, the oval sepals glabrescent, 7 — 9 mm long tapering toward base; 
petals broadly oval, 5mm long, 3.5mm broad, 3-nerved, white; stamens 
shorter than calyx; ovary half- sunken, with short styles as long as stamens; 
capsule rounded-ovoid, with very short style. June — July. 

Rocks on seashores.— Arctic: An.; Far East: Kamch. Gen. distr.: 
Ber. Described from E.Siberia. 

42. S.exilis Stephan in Sternb., Rev. Sax. Suppl. 1 (1822) 8, non Poll.; 
Engl.et Irmsch., 1. c, 267, f. 64 c-1; Kom., Fl. Kamch. II (1929) 207.- 

S. elegans Chamisso in Linnaea VI (1831) 556, non Zeyh. — S. radiata 
Small, N. Am.Fl.XII (1905) 128. -Ic: Sternb., Suppl. l,tab. 3. 

Perennial; stems numerous or solitary, erect, slender, often reddish, 
minutely glandular-pubescent, 8 — 25 cm high; radical leaves few, small, 
the blade with 3 — 7 oval, acuminate lobes, 4—7 mm long, 7 — 9 mm broad, 
with cuneate or rounded base, glabrous, the petioles 2—4 times as long as 
blade, puberulent; cauline leaves sessile, with 3 acute lobes, cuneate; 
flowers few, in axils of upper leaves, on erect or declinate, densely pubescent 
pedicels, twice as long as flowers; calyx dissected nearly to base, with 
acute, oblong, reflexed sepals, 2 — 2.5 mm long, 1 mm broad, 3-nerved; petals 
white, oblong-oval, 10 — 12 mm long, 4— 5mm broad, slightly dentate; stamens 
slightly longer than sepals; ovary slightly sunken, with short styles 1 mm 
long; capsule oblong, with strongly divergent styles. July. 



135 



174 River and stream banks. — Arctic : Arc. Sib., Chuk., An.; Gen. distr. 

Ber.,N.Am. Described from Siberia. 



Section 7. DACTYLOIDES Tausch, Hort. Canal. (182 3) p.p.- Miscaria 
Haw., Enum. Sax. (1821 ) 36-39; Small, 1. c, 120 (sub gen.).- Flowers 
campanulate, with developed hypanthium; ovary sunken, oval or rounded, 
less often flattened; petals usually white, less often yellowish, reddish 
or greenish. Above-ground cauline, scarcely leafy shoots cespitose; 
leaves usually 3— 5 -sect, less often entire. 



Cycle 1. Androsaceae Engl, et Irmsch., 1. c, 298. — Leaves oblong- 
lanceolate, spatulate, cuneate, entire; cauline shoots loosely leaved. 
Flowers solitary; petals twice as long as sepals. 

43. S.androsacea L., Sp.pl. ed. I (1753) 399; Ldb., Fl. Ross. II, 225. - 
S.androsacea var.uniflora Kryl. in A. H. P. XXI (1903) 1 6; Kryl., 
Fl.Zap. Sib. VI (l 931 ) 1420; Engl, et Irmsch., 1. c, 302, f. 71.- Ic: Sternb., 
Suppl. I, tab. XI, f. 4. 

Perennial, densely cespitose,with short loosely leaved cauline shoots, with 
leaf rosettes at the summit; flowering stems leafless or 1—2 -leaved, erect, 
2—6 cm high; radical leaves entire, oblong -lanceolate, subobtuse, tapering 
to broad petiole, 5 — 10 mm long, 1.5— 3 mm broad, 3— 5 -veined, sparsely 
glandular -hairy; cauline leaves sessile, narrowly lanceolate, smaller; 
flower solitary, terminal, 0.5—0.6 mm in diameter; calyx dissected to the 
middle, sparsely glandular, with ovate, obtuse sepals; petals white, 2—2.5 
times as long as sepals, oblong -obovate, 5.5— 6.5 mm long, 2.5— 3 mm broad, 
3 -nerved; pistils with half-inferior ovary dissected to the middle; stamens 
shorter than sepals. June. (Plate X, Figure 6). 

Rocks among stones, moist soils, moss -and -lichen tundra in the Alpine 
region.— W.Siberia: Alt.; E.Siberia: Ang. -Say. Endemic. Described 
from Siberia. Type in London. 

European forms represent a separate race, if not an independent species 
and are therefore not cited in the general distribution. 



Cycle 2. Caespitosae Engl, in Engl, und Prantl, Nat. Plfzf. Ill, 2a 
(l 891 ) 55.— Densely cespitose plants; leaves 3— 5 -sect into oval lobes; 
petals 2—3 times as long as sepals. 

44. S. sileniflora Sternb., Suppl. II (1831 ) 68, tab. 13; Ldb., Fl. Ross. II, 
224. — S. c a e s p i t o s a var.uniflora Hook, et Arn., Bot. of Beech, (l 832) 
123.- Miscaria sileniflora Small, N. Am. Fl. XXII (1905) 120. 

Perennial, densely cespitose; cauline shoots short, loosely leaved; 
flowering stem to 2.5 cm high, thickish, glandular-pubescent, sparsely 
leafy, few, often 1 -flowered; radical leaves with flat petioles, 3-, sometimes 
5 -partite, the lobes tapering upward, the lateral shorter than the middle 
lobe, all lobes rhomboid in outline, with 2—3 veins confluent into 1 broad 
vein, the blade rhomboid in outline, sparsely pubescent on both sides: cauline 



175 



136 



76 



leaves usually oblong-linear, sessile; flowers erect, to 1 cm in diameter; 
calyx dissected to 1 / 3 , oval, inflated at base, constricted at base of sepals; 
sepals oblong, obtuse; petals white, oblong-obovate, 2—3 times as long as 
sepals, 3 -nerved; ovary obovate; styles short. There is no data about 
the time of anthesis or the ecology of this species. 

Arctic: Arc. Sib., An. Gen.distr.: Ber. (Unalaska, Chamisso Island, 
Eschscholtz Bay). Described from Unalaska Island. 

Note. In the herbarium of the Biological Institute, there is one specimen from 
the herbarium of M. Bieberstein "ex Sibiria;" there is no more exact indication 
of the distribution area of this species in the Soviet Union. Ledebour reports 
this species for Lavrentiya Bay (collection of Eschscholtz); it is quite 
possible that the specimen from Bieberstein's herbarium, dated 1809, 
is the one collected by Eschscholtz and reported by Ledebour. 

45. S.lactea Turcz. in Bull. Soc. Nat. Mosc. XIII (1840) 71; Engl., Mo n. 
Sax. (1872) 92; Engl, et Irmsch., 1. c, 384. 

Perennial, loosely cespitose, with short, leafy cauline shoots; flowering 
stems erect, branching, glandular -pubescent; leaves thin, 0.8—1.5 cm long, 
cuneate, palmatisect into 3-4 upward -tapering lobes, the lower cauline 
and the radical leaves petiolate, the upper cauline sessile, shorter than 
the others; inflorescence oval-oblong, paniculate, with slender, slightly 
glandular pedicels of various length; flowers erect, less often nodding, 
campanulate, 0.6—0.8 cm in diameter; calyx dissected to the middle, with 
rounded base and oblong-triangular subobtuse sepals 1.5 mm long, slightly 
glandular; petals white, oblong-obovate, cuneate, 5—5.5 mm long, 2.5—3 mm 
broad, with slender glands; stamens not exceeding calyx; pistils with short 
styles . slightly longer than stamens. June — July. 

Riverbanks, moist soils. — E. Siberia: Lena-Kol.; Far East: Okh. Endemic. 
Described from the Okhotsk Territory after a specimen collected between 
the Chernolesskaya and Allakh railroad stations. Type in Leningrad. 

46. S.caespitosa L., Sp.pl. ed. I (1753) 404; Ldb., Fl. Ross. II, 224; 
Kryl., Fl. Zap. Sib. VI (l83l) 1419; Kom., Fl. Kamch. II (1929) 214. - 
S.caespitosa subsp. e u c a e s p it o s a Engl, et Irmsch., 1. c, 369. — 
Miscaria caespitosa Haw., Enum. Sax. (1821) 37.- ? S.groen- 
landica L., Sp.pl. ed. I (1753) 404.- Ic: Sternb., Suppl., tab. Xlb, f. 1 . 

Perennial, with slender rhizomes, densely cespitose, with branching 
cauline shoots; flowering stems erect, 2— 20 cm high, glandular -puberulent; 
radical leaves in dense rosettes, petiolate, palmatisect into 3—5 oblong- 
linear, obtuse lobes 3—12 cm long, 2 — 10 cm broad; cauline leaves few, 
the lower smaller, 3 -sect, the upper entire, sessile; flowers usually 3, less 
often 1 (f. uni flora (R.Br.) Engl.), terminal, erect, broadly campanulate, 
0.6— 0.7 cm in diameter; calyx densely glandular, tapering downward, 
dissected to x / 3 into triangular ovate, obtuse sepals; petals obtuse, 3 times 
as long as sepals, 3 -nerved; stamens shorter than petals but longer than 
calyx; pistil divided to l / 3 into styles, with half-inferior ovary; capsule 
adnate to 2 / 3 to calyx, to 10 mm long. July. (Plate X, Figure 7). 

Forest and Arctic zones, stony slopes, tundra. — Arctic: Nov. Z., Arc. 
Sib.,Chuk.,An.; European part : Kar. -Lap., Dv.-Pech., N. Urals; W.Siberia: 
Ob. (N. part); E.Siberia: Yenis. (N. part), Lena-Kol., Ang. -Say. Gen.distr.: 
Arc, Scand., Arc. Am. Described from Lapland. Type in London. 



137 



177 



Note. Engler and Irmscher record subsp. d e c i p i e n s (Ehrh. ) Engl, 
et Irmsch. with many varieties and forms for W. Europe; these differ 
so strongly from the Arctic forms that they should be regarded as 
independent species, viz., S. d e c i p i e ns Ehrh. and S.bohemica Panzer. 
There is some doubt as to the identity ofS. caespitosa L. with 
S.groenlandica L. Engler and Irmscher identify S. groenlandica 
partly with S. caespitosa subsp. decipiens Ehrh., and partly with 
subsp. eucaespitosa and f. uniflora. Litvinov, however, classes 
the herbarium part of the Siberian material as S.groenlandica. 
These definitions characteristically correspond to low-growing specimens. 
In this particular case, the size of the plant cannot be considered as a 
systematic character for the differentiation of 2 species, since all the 
more southerly forms are larger within the limits of the described species. 
It is very difficult to ascertain the true status of S. groenlandica L. 
owing to the variety of the material from Greenland. 



Cycle 3. Exarato-moschatae Engl, et Irmsch., 1. c, 388. — 
Very densely cespitose. Leaves 3— 5 -sect; lobes obtuse, linear, with 
prominent nerves. Cauline shoots densely leafy, lignifying; petals 
1.5—2.5 times as long as sepals. 

47. S.terektensis Bge. in Ldb., Fl. Alt. II (1830) 123. - S. mos chata var. 
terekt ensi s (Bge.) Engl, et Irmsch., 1. c.,432; Kryl., Fl. Zap. Sib. VI (l 931 ) 
1419.- S.muscoides Ldb., Fl. Ross. II, 223, non Wulf. - Ic. : Ldb., Ic. Fl. 
Ross. (1834) tab. 398. 

Perennial; tufts large, rather compact, with densely leafy shoots; 
flowering shoots erect, glandular -pubescent, 2 — 8 cm high; leaves to 12 mm 
long, incised nearly to the middle into 3 oblong -linear, upward -tapering 
lobes, 1—4 mm long, 0.5—1.5 mm broad, cuneate toward base, glandular- 
hairy; cauline leaves trilobate or entire, linear; flowers usually solitary, 
less often 2 or 3; pedicels glandular, with obtuse bracts; calyx dissected 
to 7 3 — V 2 , tapering downward, with oval-triangular, obtuse, erect sepals; 
petals twice as long as sepals, yellowish, rounded -oval, 3 -nerved, 
4.5—5 mm long, 2.5—3.5 mm broad; pistils with nearly inferior ovary 
and very short styles. July. 

Alpine zone, rocks, taluses, moraines. — W.Siberia: Alt.; E.Siberia: 
Ang. -Say.; Centr.Asia: Dzu. -Tarb. (Saur). Gen. distr. : Mong. Described 
from the Altai, from the Terektinskii Range, between the Ursul and Koksu 
rivers. Type in Paris. 

48. S.moschata Wulf. in Jacq., Misc. austr. II (1781) 128; Grossg., FL 
Kavk.II (1932) 236.- S.muscoides Wulf in Jack. Misc. austr. II (l78l) 
123, non Allioni. - S. c a e s p it o s a L., Sp.pl. ed. I (1753) 404, p. p. - 

S. exarata subsp. m o s c ha t a Caval. in Burnat, Fl. Alp. mar. V, 1 (1913) 
81 . — S. m o s c hat a subsp. e um o s c ha t a var. 1 o ng i p e t a 1 a Engl, et 
Irmsch., 1. c, 414. - Ic. : Sternb., Rev., tab. XI, f. 2. 

Perennial, loosely or densely cespitose; cauline shoots herbaceous or 
semilignified, densely leafy, with a terminal rosette of green leaves; 
flowering stems erect, 2 — 12 cm high, usually numerous, glabrous or slightly 



138 



glandular; leaves thin, smooth with inconspicuous veins, glabrous, 
minutely glandular on the margin, oblong-spatulate, cuneate, dissected 
to 2 — 3, less often 5 obtuse, linear spatules; cauline leaves smaller, with 
shorter undissected part; inflorescence 3— 9 -flowered, corymbiform; pedicels 
densely glandular, slender, with oblong, entire bracts at base; flowers 
0.5—0.8 cm in diameter, petals greenish, sometimes reddish, spreading, 
sublinear or tapering to base, obtuse with 3 parallel nerves, 0.2—0.3 mm 
long, as long as or slightly longer than calyx; calyx dissected to the 
middle into oval sepals; stamens shorter than corolla, with lilac or yellow 
anthers; capsule ovoid -subglobose. Entire plant has a strong musky 
scent, especially the leaves. June. (Plate X, Figure 9). 

Rosks, gravel, sands, alpine meadows in the Alpine zone. — Caucasus: all 
regions except W. Transc. Gen. distr. : Centr. Eur., W. and E.Med., 
Bal.-As. Min. Described from Germany. Type in Berlin. 

Note. The plant varies in overall size, density of tuft, and size of 
leaves. The following forms can be differentiated in the Caucasus: 
l)f.vulgaris Engl., Mon. Sax. (l 872) 1 79; f.intermedia (Mert. et 
Koch) Engl, et Irmsch., 1. c, 419; Oettingen in Fl. cauc. crit., f. 40 (1913) 
22 (excl. syn.). — the typical form, which occurs most often; 2) f. m i nut a 
Oetting., 1. c, 26 = S. pubescens M.B., Fl. taur. cauc. Ill (1819) 2 94, 
non Sternb. ex Poir. — a low-growing, densely cespitose, few -flowered form; 
3) f. laxa (Sternb.) Engl., Mon. Gat. Sax. 1872 (177); Oetting., 1. c, 26. - 
a loosely cespitose form with larger petiolate leaves, many -flowered. 

49. S. pontica Albow, Prodr. Fl. colch. in Tr. Tifl. Bot. Sada in 1 (1895) 
97.— S.muscoides f. compacta Mert. et Koch, Deutschl. Fl. Ill 
(1831 ) 147. — S. m o s c ha t a f. compacta Oetting. in Fl. cauc. crit. f. 40 
(1913) 24; Engl, et Irmsch., 1. c, 419, p.p. 

Perennial, densely cespitose; cauline shoots branching, imbricate- 
leaved, often columnar; flowering stems small, 2—4 cm high, very slender, 
glabrous; leaves in a rosette at apex of cauline shoots, cuneately tapering 
to petioles, divided into 3 oblong-linear apically rounded lobes, with 
3 veins not reaching the lobes and confluent in the petiole into a single vein, 
glabrous, minutely glandular on the margin; cauline leaves linear, small; 
flowers short -pediceled, gaping, 6— 7 mm in diameter; calyx dissected to 
beyond the middle, with narrow sepals, glandular; petals greenish, pink, 
or reddish, linear, 3-nerved, 3mm long, slightly longer than sepals; 
stamens with greenish anthers, shorter than petals. July. 

Limestones, gravels, and sands in the alpine zone. — Caucasus: 
W. Transc, Dag., Cisc. Endemic. Described from Mt. Adzhituko in 
Abkhazia. Type in Geneva. 

Note. Species very closely related to S. moschata, differing in its 
tufts, columnar shoots and glabrous stems. Parallel to S. adenophora 
Koch, which is as close to S. exarata Vill. as the described species 
is to S. m o s c h a t a Wulf. 

50. S. exarata Vill., Prosp.d. l'hist.d. PI. Dauph. (1779) 47; Ldb., Fl. 
Ross. II, 224; Oetting. in Fl. cauc. crit. f. 40 (1913)27; Engl, et Irmsch., 
I.e., 405 p. p. ; Grossg., Fl.Kavk. II (1930) 236.- S. nervosa M. B., Fl. 
taur. -cauc. I (1808) 314, non Lap.- S. mixta M. B., Fl. taur. -cauc. II 
(1808) 460, non Lap. 



139 



179 



Perennial, loosely cespitose; cauline shoots slender, herbaceous and 
lignifying, densely or loosely leaved; flowering stems erect, covered 
with soft hairs below, glandular above, 3—12 cm high; leaves in a rosette 
at apex of cauline shoots, petiolate or sessile, cuneate, palmatisect, 
oblong-linear, with 3, less often 5 obtuse or acuminate lobes, the 2 lateral 
lobes often bifid, all prominently veined, glandular on the margin, to 2 cm 
long; cauline leaves smaller, 3 -fid or entire, linear -lanceolate; flowers 
4—10 in a corymbiform inflorescence, to 1 cm in diameter; calyx half 
as long as corolla, dissected to the middle, with rounded base, sepals 
obtuse or slightly acuminate, oblong, glandular; petals white or slightly 
yellowish, sometimes reddish, oval, 3 -nerved to 6 mm long, 3 mm broad; 
capsule ovoid, subobtuse below. July. (Plate X, Figure 8). 

Stony and rocky soils in the alpine zone. — Caucasus: Cisc.,W.and E.Transc. 
Dag.; Gen.distr.: Centr. Eur., W. and E. Med., Bal.-As. Min., Arm. -Kurd. 
Described from Dauphine, France. 

Note. Very variable species, forming several ecological forms and 
varieties. The following forms have been reported for the Caucasus: 
1) f. vulgar is Engl., Mon. Gat. Sax. (1872) 179; Oetting. in Fl. cauc. crit. 40 
(1913) 28.- S. exarata var.compacta Koch, Syn. (1837) 274. - 
S.exarata var. villa r s ii Engl, et Irmsch. (1919) 406.— the most widely 
distributed, typical form; 2)f. laxa Koch, 1. c, Syn. (1837) 273; Oetting., 
Fl. cauc. crit. 40 (1913) 29.— cauline shoots elongated; leaves long-petioled, 
5 -sect, the lateral slightly smaller; nonflowering longer than flowering 
stems; petals yellowish white. Less widely distributed; 3) f. i nt r i c at a 
(Lap.) Oetting., 1. c.,29.- S. int r icata Lap., Fl. pyr. (1801 ) 58; Engl., 
Mon. Gat. Sax. (1872) 180; Engl, et Irmsch., I.e., 402.- S. nervosa J3 
minor Sternb., Rev. (1810) 52. — densely cespitose; stems delicate; panicle 
loose; leaves trilobate, the 2 lateral lobes bifid; petals rounded -oval. 
These forms grow under different ecological conditions in the same 
districts; f.laxa Koch is a shade -loving form, f. v u lga r i s usually 
occurs in sunny sites. 

51. S. adenophora C.Koch in Linnaea XIX (1847) 40; Boiss., Fl. or. II, 
805. - S. exarata var. or ientalis Engl., Mon. Sax. (1872) 180; Oetting. 
in Fl. cauc. crit. f. 40 (1913) 30. — S. exa rat a var. adenophora Engl. 
et Irmsch., I.e., 414.- Exs. : Sintenis, Iter or. (1892), No. 4739 (1891 ), 
No. 7055. 

Perennial, densely cespitose; cauline shoots long, branching, densely 
covered with imbricate old leaves, with a rosette of young green leaves 
at the apex; leaves thickish, compactly overlapping, rounded toward base, 
palmatisect into 5 obtuse lobes, very prominently veined, to 12 mm long, the 
lobes 2—4 mm long, glabrous, sparsely glandular on the margin; flowering 
stems slender, erect, glandular; inflorescence crowded, few-flowered; 
pedicels short; flowers 6— 8 mm in diameter; calyx dissected to the middle, 
1 „ with obtuse, oval sepals, glandular; petals twice as long as calyx, white, 
obovate, 3 -nerved, 5 mm long, 3 mm broad; stamens shorter than petals; 
capsule globose. July. 

Alpine meadows.— Caucasus: W., S., and E. Transc. Gen.distr.: 
Arm. -Kurd., Bal.-As. Min. Described from Asia Minor. Type in Berlin. 



5773 140 




PLATE X. 1-Saxifraga bronchialis L. , a) leaf; 2-S.spinulosa Adams, a) leaf; 3 - 
S.anad y rensis A. Los., a) leaf; 4 - S. f irma Litv.,a) leaf; 5 - S. che r le ri o id e s D.Don., a) leaf; 
6 — S.androsaceaL.; 7-S.caespitosaL.,a)leaf; 8 — S. e x a r a t a Vill., a) leaf; 9-S.moschata 
Wulf, a) leaf. 



141 



52 . S. verticillata A. Los. sp. nova in Addenda VIII, p. 3 73.— S. exarata 
var. int ricata Som. et Lev., A. H. P. XVI (1900) 173, non Lap. - 
S. exarata var. kusnezowii Engl, et Irmsch., 1. c, 412. — Ic. : Engl, 
et Irmsch., 1. c., f. 94. F. 

Perennial, loosely cespitose; cauline shoots short; flowering stems 
numerous, profusely branching in inflorescence from middle of stem, 
5 — 15cm high, strongly glandular; leaves in whorls, spatulate, 1.5 cm long, 
short -petioled, 3— 5 -partite, lobes short, obtuse, with recurved, cochlear 
tips, dark green, with very prominent veins, numerous below and reaching 
base as a parallel cluster, usually 2 nerves per lobe; cauline leaves 
smaller, trilobate; inflorescence loosely corymbiform-paniculate; pedicels 
strongly glandular, of various length, slender; flowers 0.6— 0.8 cm in 
diameter; calyx with rounded base, inflated in fruit, dissected to the 
middle, with oblong, slightly acuminate sepals, about half as long as 
petals, strongly glandular; petals white, oval, 3—4 mm long, 3 -nerved; 
stamens shorter than petals, with spherical white anthers; pistils with 
divergent short styles; capsule rounded -ovoid. June. 

Pine forests of the upper forest zone, taluses. — Caucasus: Cisc, 
Endemic. Described from the Klukhori Pass. Type in Leningrad. 

Note. Very closely related to S. ex a r at a Vill.; the loose leaf 
whorls and the branching inflorescence are characteristic. 

Section 8. TRACHYPHYLLUM Gaudin, Fl. Helv. Ill (1828) 85, 108.- 
Ciliaria Haw., Enum. Sax. (l 821 ) 41 et Leptasea Haw., 1. c, 40 (sub 
gen.). — Ovary sunken; flowers broadly campanulate; styles curved, 
short; cauline shoots densely leafy, densely cespitose; leaves narrow, 
hispid, glandular, mucronate. 

18 _ 53. S. anadyrensis A. Los. sp. nova in Addenda VIII, p. 373. 

Perennial, loosely cespitose; cauline shoots elongated, covered with 
erect leaves; flowering stems reddish, glandular, sparsely leafy, 8—12 cm 
high; leaves bright green, with dark violet-red apex, the old leaves 
violet or light brown, sharply keeled, concave -canaliculate above, strongly 
acuminate, with white mucros, short -glandular at the margin, 6— 8 mm 
long, 0.8 — 1 mm broad; cauline leaves linear, acute, slightly shorter, few; 
inflorescence corymbiform-paniculate; pedicels glandular, 3 — 10 mm long; 
bracts subobtuse, linear; calyx dissected nearly to base, sepals narrowly 
triangular, bluntly acuminate, reddish, minutely glandular on the margin; 
petals oblong -oval, tapering downward, yellowish with darker dots in lower 
part, to 5 mm long, 1 mm broad, 4 times as long as sepals; stamens shorter 
than petals; filaments dilated at base; pistils shorter than stamens, with light 
green oval ovary and erect, rather thick short styles; capsule ovoid, 
with divergent styles. July. (Plate X, Figure 3). 

Stony slopes. — Arctic: Arc. Sib., An. ; E.Siberia: Lena-Kol. Endemic. 
Described from the upper reaches of the Anadyr River, from the Medvezhi 
Mountains. Type in Leningrad. 

54. S. bronchialis L., Sp.pl. ed. I (1753) 400; Ldb., Fl. Ross. II, 207, p.p. - 
S. b r o n c hi a 1 i s var.genuina Trautv. f. p i c e i f o 1 i a Engl, et Irmsch., 
1. c, 459. 



142 



Perennial; rhizome creeping, slender; cauline shoots short, densely 
leafy; flowering shoots 10 — 35 cm high, glabrous, sparsely leafy; leaves 
remote, sessile, linear-lanceolate or subulate, to 12 mm long, to 1.5 mm 
broad, keeled below, with rigid cartilaginous white mucro to 1 mm long, and 
with crowded pectinate, acute, flat, white prickles; cauline leaves shorter, 
narrowing more abruptly toward the apex; inflorescence corymbiform- 
paniculate; pedicels glandular, with small linear bracts; flowers 8 — 12, 
to 8 mm in diameter; calyx dissected to j$, with oval -triangular sepals, 
glabrous; petals light yellow with purple dots, 3 -nerved, oblong-oval, to 
4 mm long, to 0.5 mm broad, nearly 3 times as long as calyx; stamens 
slightly shorter than petals; pistils with short styles and suborbicular ovary. 
(Plate X, Figure l). 

Sandy shore slopes, rocks, gravel.— Arctic: Arc. Sib.; E. Siberia: Yenis. 
(N. part), Lena-Kol., Dau., Ang. -Say.; Far East: Okh., Uda, Uss., Ze. -Bu., 
Sakh. Endemic, (reported for N. Am., but it is more correct to consider the 
American forms as independent species). Described from Siberia. Type in 
London. 

55. S. spinulosa Adams in Mem. Soc. Nat. Mosc V (1817) 96. — S. br on - 
chialis var. g enuina Trautv. in A. H. P. VI, 1 (l 879) 20, p. p. —S. br on - 
c h i a 1 i s subsp. spinulosa (Adams) Hulten, Fl. of Kamtsh. Ill (l 929) 
14. — S. bronchial is var. multiflora Ldb., Mem. Acad. Peter sb. V 
(1812-1814) 532. 

Perennial, loosely cespitose; shoots to 5 — 6 cm, leafy, with erect leaves; 
flowering shoots to 20 cm, glabrous, leafy, often glandular in upper part; 
leaves lanceolate or oblong -lanceolate, flat above, slightly convex below, 
lustrous, coriaceous, acuminate, to 15 mm long, to 2 mm broad, mucronate 
and ciliate, often glandular on the margin; inflorescence corymbiform- 
paniculate; pedicels glandular, of various length, with small bracts; flowers 
to 12 cm in diameter; calyx dissected to beyond the middle, with oval sepals; 
petals yellow, 3 -nerved, with purple dots, oblong-oval, 5 — 7 mm long, 
2.5 — 3.5mm broad; stamens shorter than petals; pistils with rounded 
ovaries; styles to 1 mm long; capsule oblong with divergent, elongated 
styles. July. (Plate X, Figure 2). 

Open pine forests, rocks, sandy soils, pebbles, slopes. — Arctic: Arc. 
Eur., Arc. Sib.; European part: Dv. -Pech. (N. Urals), V. -Kama., (Urals); 
W.Siberia: Ob, Alt.; E.Siberia: Ang. -Say., Dau., Lena-Kol.; Far East: 
Ze. -Bu., Gen. distr.: Mong., Ch. Described from Kamchatka. Type in 
Leningrad. 

Note.l. Litvinov reports for Sakhalin var. pseudoburseriana 
Schmidt with broader and more lustrous leaves; it deserves separation 
as an independent species, but lack of material renders this impossible. 

Note 2. S. bronchialis L. and S. spinulosa Adams are two very 
closely related forms. Abundant material from Siberia shows that there 
are many transitional forms, which apparently hybridize strongly. The 
permanent distinguishing characters are flower size and leaf shape. 
S. spinulosa Adams varies greatly; the Arctic and bald -mountain (Sayans) 
forms are smaller and more compact, with denser, somewhat curved but 
always flat leaves; the Ural and Russian-European are distinguished from 
the typical (Transbaikalian) forms by their larger flowers, but cannot be 
classed as S. bronchialis owing to leaf shape. 



143 



56. S. firma Litw. sp. nov. in sched. (descr. in Addenda VIII, p. 374. — 

S. bronchial. is Kom., Fl. Kamtsch. II (l 92 9) 217, p. p. —S. bronchialis 
subsp. spinulosa Hulten, Fl. of Kamtsh. Ill (1929) 14 p. p. 488. 

Perennial; loosely cespitose, dense; cauline shoots elongated, loosely 
leaved; flowering shoots 4 — 10cm long, slightly glandular, sparsely leafy; 
„ r leaves bright green, sometimes reddish, with concave tips of one leaf 

bending over the next, forming rosettes at tips of shoots; leaves lanceolate, 
lustrous, mucronulate, ciliate-margined, 6 — 8 mm long, 1 mm broad, the 
cauline leaves smaller and narrower; flowers in a corymbiform 
inflorescence, 3 — 8; pedicels glandular, 0.4 — 0.8 mm long, with small bracts; 
calyx glabrous, dissected to the middle, with oval sepals; petals oval- 
lanceolate, yellow, 3 -nerved, without purple dots, 5 — 6 mm long, 1 mm broad, 
3 — 4 times as long as sepals; stamens longer than petals; pistils with 
obovate ovary and rather long styles, reflexed at maturity, with oval stigmas; 
capsule compressed oblong-ovoid. July. (Plate X, Figure 4). 

Pebbles, shores, rocks. — Arctic: Chuk.,An.; E.Siberia: Lena-Kol.; 
Far East: Kamch., Sakh. Gen. distr. : Ber (?). Described from the 
Chukchi Peninsula and from Kamchatka. Type in Leningrad. 

Note. Forms transitional between S. spinulosa Adams and S. firma 
Litw. occur in Kamchatka and Yakutia, and forms transitional between 
S. firma Litw. and S. cherlerioides D. Don in Kamchatka. 

57. S. cherlerioides D. Don in Transact. Linn. Soc. XIII (1822) 382; 
Kom., Fl. Kamch. II (1929) 217. — S. bronchialis var. cherlerioides 
Engl. Mon. Sax. (1872) 216. - S. nitida Ldb. Fl. Ross. II (1844) 207.- 

S. bronchialis subsp. Funstonii Hulten, Fl. Kamtsh. Ill (l 929). — 
Ic. : Kom., Opred. rast. Dal'nevost. kraya, tab. 1 82, f. 2 — 4. 

Perennial; tufts compact, dense, with creeping rhizomes; cauline shoots 
short, imbricately and densely leafy; stems 5 — 10 cm high, slightly glandular, 
sparsely leafy; leaf rosettes crowded at tips of shoots; leaves oval- 
elliptic, thickish-concave, bluntly acuminate, ciliolate margined, 2 — 4 mm 
long, 1.5 — 2 mm broad, the cauline leaves oblong, flat; entire plant often 
reddish, inflorescence corymbiform, 2 — 8-flowered; flowers to 1 cm in 
diameter; sepals oval-triangular, glabrous; petals 2.5 times as long as 
sepals, oblong-oval, tapering upward, 3 -nerved, white or yellowish-pinkish; 
stamens shorter than petals, with rounded ovary and short styles. July. 
(Plate X, Figure 5). 

Rocks, taluses. — Arctic: Arc. Sib., Chuk., An.; E.Siberia: Lena-Kol.; 
Far East: Kamch., Ze. -Bu., Okh. Described from Kamchatka. 

Mote. For Kamchatka V. L. Komarov separates f. elongata with 
1 -flowered stem, less densely leafy, with elongated, branching cauline shoots. 

58. S. eschscholzii Sternb., Rev. Suppl. I (1822) 9, tab. 10, f. 2; Engl., 
Mon. Sax. (1872) 212; Engl, et Irmsch., 1. c, 764; Kom., Fl. Kamch, II, 
303.— S. f imbr iata D. Don. in Transact. Linn. Soc. XIII (1822) 410. 

^g Perennial, forming subspherical tufts, with densely imbricate-leaved 
stems; leaves obovate, tapering downward, strongly concave, ciliate- 
margined, membranous, 4 mm long, straw-colored or silver; flowers with 
very short pedicels; calyx with ovate, fimbriate sepals recurved after 



144 



anthesis, 1—1.5 mm long; petals oblong, long-clawed, 1 mm long; stamens 
as long as sepals; capsule subglobose. August. 

Rocks. —Arctic: Arc. Sib., An. Gen. distr. : Ber. Described from the 
shores of Lavrentiya Bay. 

Section 9. XANTHIZOON Griseb., Spicil. Fl. rumel. I (1843) 333.- 
Leptasea Haw., Enum. Sax. (1821) 39 p. p. (sub. gen.).- Ovary sunken; 
flowers gaping; petals yellow; leaves with 1 pit at the apex, rather 
sparsely covering perennial.loosely cespitose cauline shoots. 

59. S. aizoides L. Sp. pi. ed. I (1753) 403; Engl, et Irmsch. 1. c, 446; 
Kryl., Fl. Zap. Sib. VI (1931 ) 1427. — S. a u t u m n a 1 i s L. in Jacq. Enum. 
Horti Vindob. I (1762) 21. — Ic: Sternb. Rev. Sax. tab. 7, f. 1. 

Perennial; rhizome slender, creeping, loosely cespitose; stems 
ascending, spreading, elongated, branching, densely leafy, 5 — 17 cm high, 
covered with rigid short hairs, the hairs glandular in upper part; leaves 
oblong-oval or linear, cuneate, thickish and rigid, flat above, convex below, 
glabrous or rigidly ciliate-margined, 7 — 20 mm long, 1.3 mm broad; 
inflorescence 2 — 8-flowered, paniculate; pedicels 5 — 15 mm long; calyx 
dissected to the middle, with divergent, ovate-triangular, obtuse sepals, 
the calyx tube densely glandular; petals yellow, sometimes with orange dots, 
oblong -elliptic, 4.5 — 6 mm long, 1.2 — 2.5 mm broad, about 1.5 times as long 
as sepals; stamens as long as petals; pistils parted to the middle into 
styles; capsule ca. 6 mm long, 4 mm broad. July. 

Arctic region, tundra, stony and sandy soils, seashores. —Arctic: Arc. 
Eur., Nov. Z., Arc. Sib. (Kara Tundra); European part: Kar. -Lap., Dv. -Pech. 
Gen. distr.: Arc. Eur., N. Am. Described from Lapland. Type in London. 

Section 10. EUAIZOONIA (Schott) Engl., Mon. Sax. (1872) 223.- Schott 
in Schott, Nyman et Kotschy, An. bot. (l 854) 20 (subsectio). — Chondrosea 
Haw., Enum. Sax. (l 821 ) 10 p. p. (pro genere). Ovary sunken; flowers 
gaping; leaves fleshy, coriaceous, dentate, secreting lime along the whole 
margin; pitless. 

60. S. cartilaginea Willd. ex Sternb., Rev. (1810) 5, tab. 3c; Ldb., Fl. 
Ross. II, 205; Boiss., Fl. Or. II (1872) 800; Grossg., Fl. Kavk. II, 237.- 
S.aizoon subsp. cartilaginea var. e u c a r t i 1 ag in e a Engl, et 
Irmsch., 1. c, 513.- S. cotyledon M. B., Fl. taur. -cauc I (1808) 13. 

Perennial, loosely cespitose; rhizomes slender; flowering stems 
10 — 30 cm high, erect, smooth, with sparse leaves, sparsely glandular; leaves 
in dense rosettes, oblong-liguliform or oblong-spatulate, slightly fleshy, 
coriaceous, gray-green, 1—3 cm long, 3 — 6 mm broad, cartilaginous on the 
margin, crenate-dentate, secreting lime along the whole margin, abruptly 
narrowed above, muticous, the upper part of leaf the broadest; cauline 
leaves small and narrow; leaves at the very base ciliate-margined, 2 — 3 cm 
long; flowers to 13 mm in diameter; calyx glandular, dissected to the middle, 
with oval-triangular sepals; sepals 1.5 mm long, 3 -nerved; petals white, 
to 8 mm long, 4 mm broad, broadly oval, obtuse, 3 -nerved; stamens as long 
as sepals; filaments dilated toward base; pistils shorter than stamens. 
July — August. (Plate XI, Figure 10). 



145 



Subalpine and alpine zones, rocks, limestones.— Caucasus: Cisc, Dag., 
E., W., and S. Transc, Tal. Gen. distr.: Iran. (?). Described from the 
Caucasus. Type in Berlin. 

Note. F. minor Boiss., distinguished by its smaller size, has been 
reported for the Main Range of the Caucasus; Engl, et Irmsch. report 
f.major, and f. vulgaris is noted for the Oettingen herbarium. This 
species varies greatly in leaf size and shape; separation of the above forms 
is very formal; exact determination of taxonomic units within S.carti- 
laginea sensu lato requires the study of differences at various altitudes 
under natural conditions. 

61. S. kolenatiana Rgl. in Ind. Sem. H. Petr. (1865) 3 9; Gorssg., Fl. Kavk. 
II (l 930) 237. — S. a i z o o n subsp. c a r t i la g ine a var. Kol e n a t i an a 
Engl, et Irmsch., 1. c, 513. 

Perennial; loosely cespitose; rhizomes lignifying; stems erect or 
slightly ascending, smooth, glandular, often red- violet, 2 — 40 cm high; leaves 
in loose rosettes, oblong-lanceolate, slightly tapering downward, gradually 
acuminate and mucronulate, somewhat fleshy, coriaceous, green, cartilaginous - 
crenate -dentate (in lower part teeth passing into longish cilia), 2^3.5 cm 
long, 5 — 9 mm broad, the cauline leaves smaller; inflorescence paniculate, 
the lateral branchlets to 3 cm long, with small acute bracts, 2 — 3 -flowered, 
strongly glandular; flowers to 1 cm in diameter; calyx as in preceding 
species; petals pink or purple, broadly oval, to 7 mm long, 3 -nerved; 
stamens slightly longer than sepals. July. (Plate XI, Figure ll). 

Alpine and subalpine zones, rocks, stony slopes. — Caucasus: Cisc, W., 
E., and S. Transc, Dag. Gen. distr.: Iran (?). Described from Kazbek. 
Type in Leningrad. 



Section 11. KABSCHIA Engl, in Linnaea XXXV (1867) 16.— Ovary deeply 
sunken; flowers campanulate or open; cauline shoots perennial, densely 
covered with coriaceous, rigid leaves; leaves with several lime-secreting 
pits on margin. 

62. S. juniperifolia Adams in Web. et Mohr, Beitr. z. Naturk. I (1805) 
53; Ldb., Fl. Ross. II, 206; Boiss., Fl. Or. II (l 872) 803; Oetting. in Fl. cauc. 
crit. fasc. 42, 40.— S. juniper ina M. B., Fl. taur. -cauc. I (1808) 314.— 
Chondrosea juniperina Haw., Enum. Sax. (l82l) 1 5. — S. j un i p e r i - 
folia var. of typica Engl, et Irmsch., 1. c, 549. — Ic : Sternb., Rev. Sax. 
(1810) tab.X. 

Perennial; densely cespitose; cauline shoots semi-lignified, erect, 
imbricate-leaved; flowering stems 2—7 cm long, erect, leafy, pubescent; 
leaves of cauline shoots dark green, the old ones light brown, in dense 
rosettes, often somewhat whorled -remote, rigid, spiny, lanceolate or oblong - 
lanceoJate, keeled, with 2 grooves above, glabrous on the margin, ciliolate 
only in lower part, mucronate, horizontally divergent or recurved, 5 — 18 mm 
long, 1—2 mm broad, 5 — 7 -pitted; cauline leaves lanceolate or oblong-oval, 
tapering to a somewhat fleshy petiole, mucronate, smaller; inflorescence 
racemose, ovoid; pedicels pubescent, 2 — 3 mm long, in axils of rigid, 
mucronate, ciliate-margined, oval-lanceolate bracts; sepals keeled, oblong- 
oval, ciliate-margined; petals linear-spatulate or oblong -spatulate, 



146 



3—4.5 mm long, 1 .5 mm broad, twice as long as calyx, rigid, 1 — 3 -nerved; 
stamens longer than petals; styles 3— 4 mm long, erect. June — July. 
(Plate XI, Figure 7). 

Shady sites on rocks and limestones in the alpine zone.— Caucasus: 
E. and W. Transc, Dag. Endemic. Described from "Iberia." Type in 
Leningrad. 

Note. Var. im b r i c a ta (Rupr. in Sched.) Oetting., 1. c, 44 is an 
alpine form with shortened, compact, columnar shoots and dense, appressed, 
small leaves. Oettingen also separates f. distans with whorled leaves 
and f. cinerea with small leaves; these forms are apparently ecological 
variants. 

63. S. subverticillata Boiss., Fl.Or.II (1872) 802; Oettingen, 1. c, 40; 
Engl, et Irmsch., 1. c, 553. 

Perennial, loosely cespitose; cauline shoots elongated, slender, brittle, 
slightly geniculate and nodding, ribbed and canaliculate, reddish, glabrous, 
with remote whorls of rather numerous, horizontally divergent leaves; 
flowering stems 4—5 cm high, glabrous, smooth, with sparse, smaller 
leaves sometimes ciliate in lower part; leaves linear or linear-lanceolate, 
mucronate, fleshy, strongly keeled, with reflexed margins and a central 
groove on upper surface, glabrous, to 2 cm long, 1—2 mm broad, 7— 9 -pitted; 
inflorescence paniculate -corymbiform, 5— 7 -flowered; pedicels slender, 
glabrous, 1.5—2 cm long; calyx glabrous, with oblong-ovate sepals; petals 
yellow, oblong, longer than calyx, tapering, with claw to 3 mm long; stamens 
longer than petals; capsule subglobose, with long styles exceeding stamens. 
June — July. (Plate XI, Figure 8). 

Wet rocks and limestones in the alpine zone.— Caucasus: Dag., E. Transc. 
Endemic. Described from Khevsuretiya and Dagestan. Type in Leningrad. 

64. S. colchica Albow, Prodr. Fl. colch. in Tp. Tr. Tifl. Bot. Sada 1 (1895) 
96; Engl, et Irmsch., 1. c, 555. — S. subverticillata var. colchica 
Oettingen, 1. c, 50. 

Perennial, loosely cespitose; cauline shoots elongated, covered with 
remote leave whorls, imbricate -leaved in the interstices, reddish, 
glabrous; flowering stem to 4 cm high, glabrous, leafy, the leaves 
oblong, flat, ciliate, acuminate; whorled leaves acicular, with a groove above, 
subobtuse, mucronulate, 5— 7 -pitted, ciliate in lower part, smooth, 1 cm 
long, erect, the entire whorl appressed to shoot; imbricate leaves small, 
thin, acute; inflorescence corymbiform, 3— 7 -flowered; pedicels to 1.5 cm 
long, with several bracts similar to cauline leaves; calyx with oval glabrous 
sepals, 3 -nerved, 3—4 mm long; petals oval, tapering to claw, subobtuse, 
5 -nerved, 7— 8 mm long, 4— 5 mm broad; stamens as long as petals. 
June -July. (Plate XI, Figure 9). 

Caucasus: W. Transc. Endemic. Described from Mt. Chita -Gvala. 
in Mingrelia. Type in Leningrad. 

65. S. laevis M. B., Fl. taur. -cauc. I (1808) 314, III (1819) 29 sensu 
angustiore; Ldb., FL Ross. II, 205, p.p.; Oetting., 1. c, 38. — S. laevis 
var. eulaevis Engl, et Irmsch., 1. c, 556, f. D. 



147 



Perennial, loosely cespitose; cauline shoots lignifying, rather long, 
densely imbricate -leaved in upper part; flowering stems 2—4 cm long, 
leafy, glandular -hairy; leaves of cauline shoot ligulate, concave, rigid, 
1 „_ mucronate, glabrous, cartilaginous on the margin, denticulate or ciliate 
in lower half, 5 — 7 -pitted, light green, sometimes reddish below, 4—7 mm 
long, 2—2.5 mm broad; cauline leaves oboval, mucronate, 2—3 mm long, 
light green or reddish; inflorescence corymbiform, 4— 8 -flowered; pedicels 
short, short -hairy; flowers erect; calyx dissected to beyond the middle, 
with reddish, glabrous, ciliate -margined, oval, obtuse sepals, 2.5— 3 mm long, 
1.3— 1.8 mm broad, with parallel nerves; petals oblong-obovate, 3 -nerved, 
shorter than stamens, about twice as long as sepals; pistils longer than 
petals; capsule subglobose, with long divergent styles. July — August. 
(Plate XI, Figure 4). 

Rocks, taluses, and limestones in the alpine zone.— Caucasus: Dag., 
E. Transc. Endemic. Described from the Caucasus. Type in Leningrad. 

66. S. pseudolaevis Oetting. in Acta H. Bot. Juriev. X (±910) 15; 

Ej. in Fl. cauc. crit. f. 42 (1915) 38.— S. laevis var. pseudolaevis 
Engl, et Irmsch., 1. c, 556, f. C. 

Perennial, loosely cespitose; cauline shoots elongated, strongly 
lignifying, loosely leaved with old leaves, bearing broad, rather sparse 
rosettes of live leaves; flowering stems ribbed, hairy, mainly in upper 
part, 2—5 cm high, leafy; rosette leaves oblong -spatulate, nearly flat, 
slightly keeled below, conspicuously longitudinally veined, lustrous, slightly 
fleshy, mucronate, cartilaginous on the margin, with small prickles, 
5— 9 -pitted, 5—7 mm long, 2—4 mm broad; cauline leaves appressed to 
stem, oblong-obovate, 4—10 mm long, 3—4 mm broad, mucronate, ciliate in 
lower part; inflorescence oval, paniculate -capitate, 5— 6 -flowered; pedicels 
short, hairy; calyx glabrous, dissected to the middle, the sepals broadly 
oval, sometimes slightly dentate, with conspicuous nerves converging toward 
the apex; petals twice as long as sepals, obovate, often emarginate, narrowing 
to conspicuous claw, yellow, 3.5—4.5 mm long, 5 -nerved; stamens slightly 
longer than petals; ovary subspherical, with long divergent styles. 
July— August. (Plate XI, Figure 5). 

Limestone and stony soils in the alpine zone.— Caucasus: Cisc. 
(El'brus -Kazbek region). Endemic. Described from "Iberia." Type in 
Leningrad. 

67. S. scleropoda Somm. et Lev. in A. H. P. XIII, 2 (1894)186; Oettingen, 
1. c, 46; Engl, et Irmsch., l.c.,556.— S. juniperifolia var. br a chy - 
phylla Boiss. Fl. Or. II (1872) 803.- Ic: Somm. et Lev. in A. H. P. XVI 
(1900) tab. XVIII.- Exs.: Somm. et Lev., Iter, cauc, No. 496. 

Perennial, loosely cespitose, with vertical columnar, woody cauline 
shoots densely covered with reclinate or horizontally spreading leaves; 
flowering stem to 5 cm high, hairy; leaves imbricate, linear -ligulate, flat 
. _j or slightly keeled, bluntly acuminate, mucronulate, cartilaginous on the 

margin, ciliate -dentate, with 1 upper pit (rarely several pits on the margin), 
7—8 mm long, 1—1.5 mm broad, the upper leaves light green, the old lower 
leaves bright rufous, the cauline colorless, acute, petiolate, appressed to 
stem; inflorescence paniculate, ovoid; pedicels 2— 3 mm long, in axil of 



148 



bract, with bracteole at base or mid -length of calyx; calyx ovoid -globose, 
dissected to the middle-, 10 -nerved, the sepals 2 mm long, light green, oval, 
ciliate- margined; petals oblong-cuneate, yellowish, 3—5 mm long, 1 -nerved; 
stamens slightly longer than petals; ovary with long, divergent styles. 
June. 

Alpine zone rocks. Caucasus: E. and W. Transc, Cisc. (Teberda). 
Endemic. Described from the Teberda River. Type in Leningrad. 

Note. Var. nivalis Somm. et Lev., 1. c, (var . S o m m i e r i Engl, 
et Irmsch., 1. c, 557) is densely cespitose, with smaller leaves covering 
more densely the cauline shoots. Sommier et Levier report that 
S. scleropoda also produces a form with whorled leaves which 
Oettingen separates as a variety, writing that it occurs more often than 
the typical form; lack of material and observations concerning variation 
under various ecological conditions make it impossible to draw any 
conclusions. 

68. S.abchasica Oetting. in Acta H.Bot.Juriev. VIII (1907) 88; Ej. in 
Fl. cauc. crit. f. 42 (1915) 45.— S. scleropoda var. abchasica (Oett.) 
Engl, et Irmsch., 1. c, 557. 

Perennial, densely cespitose; short lignifying cauline shoots densely 
covered with imbricate leaves; flowering stem glandular-pubescent, 
ca 4—5 cm long, leafy; leaves in approximate whorls, glaucous, oblong-oval, 
fleshy, keeled, with thin, cartilaginous margins, mucronate, with small 
prickles in lower half, 1 - or 3— 5 -pitted, 0.5-0.6 mm long, 1 mm broad; 
cauline leaves small; inflorescence oval, paniculate; calyx campanulate, 
glandular, with oval sepals; petals more than twice as long as calyx, 
sublinear, 2 (?)-nerved, 4 mm long; stamens as long as petals; styles 
elongated. July— August. (Plate XI, Figure 3). 

Limestone rocks.— Caucasus: W. Transc. (Abkhazia). Endemic. 
Described from Mt. Shmek in Abkhazia. Type in Leningrad. 

69. S. caucasica Somm. et Lev. in A. H. P. XIII, 2 (1894)188; Oettingen, 
I.e., 38; Engl, et Irmsch., 1. c, 552.- Ic: Somm. et Lev. in A. H. P. XVI 
(1900) tab. XVII. 

Perennial; tuft flat; cauline shoots erect, densely covered with 
imbricate leaves; flowering stems 2.5—3 cm high, glabrous, leafy; leaves 
on cauline shoot erect, not firm, flat above, keeled below, narrowly lanceolate, 
mucronate, tapering downward, cartilaginous on the margin, smooth, 
sometimes scabrous or short -dentate in lower part, sometimes 2— 4 -pitted 
on the margin, 5—9 mm long, 1—2 mm broad; cauline leaves elliptic, acute 
or ligulate, yellowish, often reddish, ciliate -dentate; inflorescence a 
5— 7 -flowered corymb; calyx glabrous, yellowish, with erect, oval, acuminate 
sepals, 1.5—2 mm long, 1.5 mm broad, rarely mucronate; petals cuneate, ■ 
obovate, long-clawed, 2.5—3 mm long, 0.8—1 mm broad, yellow, 3 -, less often 
1 -nerved; stamens 4—5 mm long; pistils as long as stamens, dissected to 
the middle into styles. August— September. 

Mountain passes, slopes. — Caucasus: Cisc, W. Transc. (El'brus region). 
Endemic. Described from the Klukhori Pass. 

70. S.desoulavyi Oetting. in Acta H.Bot.Juriev. X (1910) 16; Ej. in Fl. 
cauc. crit. f. 42 (1915) 47. 



149 



!r; 



Perennial, densely ecspifose; caul in< shoot :; densely leafy, with 

Inconspicuous vhorls; flowering stem Leafy,with ion;.', white hairs; leaves 

convex below, With 2 Lighl green groove;: above, oblong lanceolate, acuminate, 
mucronate, Cartilaginous Ciliate on the iria rgin below, seabi ous above, 

4 6 mm Long, 5 7 pitted; cauline Leaves narrower, tapering to petiole; 

inflorescence paniculate, narrowly ovoid; pod icols ciliate, to II inin Long, 

i . '.i narrowly Lanceolate, mucronate, ion// eiiiate bracts; sepals 
oval, ciliate, 3 nerved, converging above, i pitted; petals oblong oval, 

long -clawed, 1 nerved, yellow, 4 a mm Ion/',, 1 .a lit/iff-; as Ion/', as .sepals; 

stamens slightly longer than petals; ovary orbicular, with long, creel 
styles Longer than stamens. June July. 

Taluses, glacial moraines. Caucasus! Cisc. (Balkarlya), E. Transc. 
(S.Ossetia). Kndemic. Described from ivit. Shtulu In Balkarlya. 

71. S. kusnezowiana Getting, in Acta n. Bot. Juriev. x (1910)15; 

l.j.in Fl, cauC. crlt. f. 42 (l !H '■>) '12. S.j unipcri folia var. 
kusnezowiana (netting.) Kngl. ot lrmsch., 1. c, 552. 

I'erennial; luff gray green, many Stemmed, with Semlllgni lied imbricate 
leaved shoots; leaves oblong lam eolato, mucronate, ciliate margined only 
in Lower part, 5 7 pitted; flowering stem many -flowered; pedicels 

glabrous; calyx with glabrous, ova I, ci I iafe sepals; petals obova I, tapering 
tO Claw, 5 nerved (3, the tWO marginal nerves branching), as long as 
StamenH, twice as Ion/', as sepal:;; .styles erect, as long as stamens. July. 

Caucasus; W. Transc. Endemh . Described from the Mamison Pass. 

Note. | have no material concerning this species at my disposa I, nor 
hav I seen the type. Described by Oeffingcn after a single specimen. 

72. S. carlnatft Oettlng. In A. II. Hot. Juriev. VIII (ll)0H)«MJ; Kj.in Kl. 
cauc.crlt.42 (1915)48; Engl, et Lrmsch., 1. c, 557, 

Perennial, eoHpi lose; shoots to I a cm long, bra aching, d ens ely covered 
with leaf whorls (?), 4 .5 mm long, I mm broad, erect or arched lipcurved OT 
nodding; I eaves I i nea r, acuminate, mucronate, ciliate -margined to the 
middle, with a single pj| al the apex, a I ways keeled; flowering Stem 
I flowered; calyx glabrous, sepals oblong; petals H mm long, 2.5mm 
broad, oblong oval, tapering downward, a bo u I as long as sl.n mens, yellow. 

July. 

Hocks. Caucasus: ( ' Jsc. (lialka riya ). Kndemic. Described from 
Ml.. Shtulu in Halkariya. 

Nul e. | have seen no herbarium maternal concerning this species. 

+ T.\. S. kotsehyi lioiss., Diagn. pi. nov., ser. 2, 1 1 (l JiMi) fi.a ; Kl.Or. I] 
(1872)804; Engl, et lrmsch., 1. c, 559; Oettlngen, 1. c.,41. [c: Bot. 
Mag. tab. fiOfia. Kxs.: Siohi , Hot. Itoise, No. 332; Manissadjian, PI. or., 
No. 829; Bornmuller, PI, exsic. Anatol. or., No. i nr>4. 

Perennial, densely cespltose; shoots short, densely I mbricate -leaved; 
flowering stems i B cm high, ribbed, glandular pubescent, leafy; leaves 
on shoo is corla< eous, I eeled, with small groove above, oblong -oval, slightly 

tapering downward, acuminate, glaucous, the old leaves bla< I isli, glabrous, 
smooth, cartilaginous on the margin, 7 !) nerved, ci I jo late in lower part, 
I 8mm long,:-! 3mm broad; cauline have;; spatulate or cochlea i . 



150 



tapering to petiole, smaller; Inflorescence corymbiform paniculate; 
pedicels broadening upward, of various Length, from 2 3 mm (the upper) 
to L.5 cm (the lower), densely glandular, Ln axils of oblong Linear bracts; 
calyx with narrowly triangular sepals, L.5 2,5mm Long, glabrous or else 
glandular Ln lower part; petals oblong, cuneate, 3 5mm Long, 2 2.5mm 
broad, 3 nerved, yellow; stamens slightly Longer than petals; ovary oval, 
glandular, wi1 h ered styles, equa] to stamens. June July. 

Rocks in the alpine zone. Caucasus; possibly in W. and S. Transc 
(known from the former Kars Region). Gen. distr.: Ann. Kurd , Iran. 
Described from the Cicilian Taurus. Type Ln Geneva. 

74. S.alberti Rgl. e1 Schmalh. in A.n.r. V (1877)569; Engl. e1 Lrmsch., 
I. c, 569. 

Perennial, compactly and pulvinately cespitose; shoots reduced, columnar 
densely Lmbricate Leaved, semilignif led; flowering shoots 2 5 cm high, 
densely glandular; Leaves of cauline shoots oblong, keeled below, thickened 
a1 the ;ip''\, .'lu'i.'ii'cuii;; mid ::i r« m; 1 , . with cartilaginous margin and prominenl 
Limy bloom, 3 5 pitted, ciliate Ln lower part, 3 I nun Long, I 1 ,5 mm broad; 
upper Leaves cochlear, thinner, tapering to petiole, densely glandular; 

inflorescence a corymb of I or 5 flowers; pedicels and calyx densely 
glandular; sepals oblong ova I, ci liafe margined; petals more than twice 

.is lour, a.s eaiyx, while , 'A nerved, broadly oval, obtuse, clawed; stamen 
.i i -on i half as Long as petals, pistils with short styles as Long as stamens. 
.inly Augusl . (Plate X i. Figure 6). 

Koek crevices, near glaci ers. Cent r. Asia : T. Sh., Pam.-Al. Endemic. 
Described from i.lie Kara luira rass in i.iie Meksand rovskij Range, Type 
iii I ,eningrad, 

Noie. One specimen w Lthoul flowers, with very small, narrow 
Leaflets, very thick shoots, less densely leafy, was collected by Korzhinski, 

in the Pamir from the shore;: of Lake Kara Kill; lln;; is undoubtedly an 

Lndopondcni spe< ies, imi lack of material renders ii Impossible to give 

a complel c i\i-r.r r ipl ion. 

75. S. columnaris Schmalh. in I Jer. Deufsch. Bot. Ges.X (1892) &88, 
tab. xvii. r. u 12; Oettingen, 1. c.,48; Engl, el Lrmsch., i. c, 590. 

Perennial, Loosely and pulvinately cespitose; shoots elongated, Lignified, 

Columnar, densely imbricate leaved; flowering stems glandular, to I em 
high,! flowered, With I or 2 leaves; leaves of cauline shoots glaucous, 

subtriangular Ln section, obtusely keeled below, slightly concave above, 

With reflex apex, obtuse, filiate in lower half, 3 nun Long, I nun broad. 
5 pilled, with limy bloom; flowers erect ; calyx rlan.lular.willioblu.se, 
oval sepals, 2 nun Ion/',; petals white, many nerved, broadly oval, tapering 

downward.;; in mm lour,, I 5 mm broad: stamens shorter than petals J 
styles i mm Long; ovary subspherical. June July. (Plate XI, Figure i). 

Rocks. CaucaSUSi CISC. (Balkarlya). Kndcmic. Described from the 
vicinity of Tekhan su in Halkariya. Type Ln Leningrad. 

76. S.dinnikii Schmalh. in her. i leutsch, Bot. I fes. X (i 892) 288, 
tab. XVIII, f. 6 7; Oetting., 1. c, 48; Engl. e1 Lrmsch., 1. c, 571, 

Perennial, densely, compactly, and pulvinately cespitose; shoots short, 
densely Leafy; flowering shoots 2 4 cm high, 1 flowered, leafy, glabrous; 



I si 



(195) 




PLATE XI. 1. Saxifraga columnaris Schmalh., a) leaf, hi petal; 2-S.dinnikii Schmalh. 

a) leaf; 3 - S.a bchasica Oett., a) leaf; 4-S.laevis M.B., a) leaf, b) flower; 5 - S.pseudolaevis 

Oett., a) leaf, bl flower; 6-S.alberti Rgl.et Schmalh., a) leaf; 7 - S. j uniperifolia Adams, 

.0 leaf; 8 - S.subvertic ilia ta Oett., a) leaf; 9- S.colchica Getting, al underside of leaf; 

10 - s. carti laginea Willd, a) leaf; 11 S.kolenatiana Rgl., a) leaf. 



152 



leaves of cauline shoots glaucous, bluntly keeled below, slightly elongated 
above, thickish, linear -lanceolate, mucronate, 5 -pitted, smooth on the 
margin above, ciliate in lower part, with limy bloom, 3—6 mm long, 0.8—12 mm 
broad, recurved; cauline leaves slightly smaller, eciliate; flowers erect; 
calyx glabrous, often reddish like upper part of stem, with oval, obtuse 
sepals 2 mm long, 1 .5 mm broad; petals yellow, 3 -nerved, oblong -oval, 
twice as long as sepals; stamens as long as petals; ovary oblong with 
long styles exceeding stamens, 5— 7 mm long. June — July. (Plate XI, 
Figure 2). 

Rocks. — Caucasus: Cisc. (Balkariya). Endemic. Described from 
the vicinity of Tekan-su in Balkariya. Type in Leningrad. 



Section 12. PORPHYRION Tausch, Hort. Canal. (1823) 1.- Anthy- 
phylla Haw., Enum. Sax. (1821 ) 43 (sub gen.). Ovary sunken; flowers 
large, pink; leaves opposite, with several lime -secreting pits, densely 
covering perennial shoots. Plants forming large tufts. 

77. S.oppositifolia L., Sp.pl. ed. I (1753) 402; Ldb., Fl. Ross. II, 204 ; 
Engl, et Irmsch., I.e., 616; Kom., Fl. Kamch. II (1929) 219; Kryl., Fl. 
Sib. Zap. VI (1931) 1428.- A nthyphylla op p o s i t i f o 1 i a (L.)Four. 
in Ann. Soc. Linn. Lyon. Nouv. ser.XVI (1863) 386. — A.coerulea Haw., 
Enum. Sax. (1821 ) 43.- Ic: L., Fl. Lap. 179, tab. 2, f . 1 . 

Perennial; stems profusely branching, numerous, with ascending or 
prostrate branches, densely imbricate -leaved, densely cespitose; leaves 
opposite, oblong -obovate or subelliptic, entire, ca. 6 mm long, 2.5 mm 
broad, obtuse and thickened at the apex, 1 -, rarely 3 -pitted, flat above, 
keeled below, glabrous, ciliate-margined, basally connate in pairs, the 
leaves surrounding stem with a short sheath, decussate and closely 
approximated; flowers at ends of branches, subsessile, solitary; calyx 
dissected to 3 /i> with sparse hairs at base; sepals ovate, obtuse, ciliate- 
margined; petals pink-red (violet in herbarium), obovate, tapering to claw, 
10—11 mm long, 5—6 mm broad, 2—2.5 times as long as sepals, 5— 7 -nerved; 
stamens shorter than petals and styles; ovary almost superior, with 
long styles. July. 

Stony slopes, rock crevices, rock streams, near glaciers.— Arctic: 
Nov. Z., Arc. Sib., Chuk., An.; European part: Kar.-Lap., Dv.-Pech. (N.part); 
W.Siberia: Alt.; E.Siberia: Lena -Kol. , Dau., Ang. -Say.; Far East: 
Kamch., Okh.; Centr. Asia: Dzu. -Tarb., T. Sh. Gen. distr. : Arc. Eur., 
Mong., China, Dzu. -Kash., Him., N. Am. Described from Spitsbergen. 
Type in London. 

Note. Engler separates subsp. a s iat ica (Hayek) Engl, et Irmsch., 
1. c, distinguished by its denticular leaves. This character is not always 
maintained. 

Specimens from Pamir-Alai are different, but they cannot be separated 
owing to lack of material. 



Section 13. DIPTERA (Borkh.) Engl, et Irmsch., 1. c, 645; Borkh. in 
Roem., Neu. Mag. I (1794) 2 9 (sub gen.).— Ligularia Duwal, PI. succul. 
Hort. Alencon (1809) 11; Haw., Enum. Sax. (1821 ) 50.- Ovary free, oblong; 



153 



flowers zygomorphic, the lower 3 petals nodding, longer than the 2 upper; 
flowers always nodding; leaves fleshy. 

78. S. cortusifolia Sieb. et Zucc., Fl. jap. fam. nat. I in Acad. Munch. IV, 
2 (1843)190; Engl, et Irmsch., 1. c, 648. - Ic. : Yatabe, Iconogr. Fl. jap. I, 

1 (1891) tab. 3-6. 

Perennial; stem erect, 10— 40 cm long, leafless, glabrous or sparsely 
hairy; radical leaves rather sparsely pubescent with rigid hairs; petioles 
2—3 times as long as the blade; blade coriaceous, fleshy, rounded -reniform, 

2 — 8 cm long, 3 — 1 1 cm broad, glabrous above, hairy below, 9—1 1 -lobed, the 
lobes rounded, obtusely crenate -dentate, mucronulate; inflorescence 
corymbiform -paniculate, the branches long, ascending at an acute angle; 
pedicels with small bracts, slender, glandular, 0.5 — 1.5 cm long; flowers 
zygomorphic; calyx with triangular, spreading sepals, 1.2—1.3 mm long; 
petals varied, the lower 2 nodding, linear -lanceolate, 1—2 mm long, 
1.5—2 mm broad, the 3 upper subelliptic or oblong -oval, twice as long 

as sepals, subobtuse, short -clawed, all white or pinkish; stamens shorter 
than petals; pistils with long erect styles; capsule oblong-ovoid. 
July —September . 

Moist, shady rocks. — Far East : Uss. Gen. distr.: Jap.-Ch. Described 
from Japan. Type in Berlin. 

79. S. oblongifolia Nakai, Fl. Koreana in Journ. of College of Sc. of 
Tokyo XXVI (1909) 218, tab. 11; Engl, et Irmsch., I.e., 645. 

Perennial; stem erect, branching, sulcate, to 20 cm high, covered with 
slender glandular hairs, dense in upper part, with 1—2 leaves below 
. qq inflorescence; radical leaves petiolate, the blade oblong, oblong-oval, or 
oval, light green, reddish below, 3—7 cm long, 2—5 cm broad, subtruncate 
or cordate at base, broadly sinuate -dentate with apiculate teeth; leaves 
glabrous above, glandular -pubescent below; petioles sulcate, glandular, 
as long as or longer than the blade; inflorescence broad, spreadingly 
paniculate; pedicels slender, glandular, 3— 10 mm long; flowers small; 
calyx with triangular sepals, glabrous; petals yellow, oblong, 2 mm long; 
stamens with subulate filaments about as long as petals; pistils with short 
recurved styles; capsule 3 mm long. July. 

Rock crevices, moist, shady gorges.— Far East: Uss. Gen. distr. : 
N.Ch., Korea. Described from Korea. Type in Tokyo. 



Genus 707. MITELLA * L. 

L.Sp.pl.L. (1753) 406. 

Flowers bisexual, 5-merous; calyx campanulate, with short tube and 
5 divergent sepals; petals 5, pinnate, with thin lobes; stamens 10, with short 
filaments and bilocular, cordate anthers; pistils with half-inferior ovary 
and 2 short styles; capsule unilocular, many-seeded. Perennial herbs 
with simple leaves. 



Diminutive from the word mitra.owing to the plant's likeness to a miter. 



154 



1. M. nuda L., Sp.pl. (1753) 406; Ldb., Fl. Ross. II, 228; KryL, Fl. Zap. 
Sib. VI, 1431; Kom. and Alis.,Opred. rast. Dal'nevost. kraya I (1931) 609.- 
M. cordifolia et M. r e n i f o r mi s Lam., Encycl. Meth. IV (l 797) 1 96. - 
Piarella uniflora Retzius, Observ. bot. (l 783) 30. - Ic. : Lam., 111. 
tab. 373, f. 2-3; Britton and Brown, Fl. Un. St. and Can. II, 181; Nekrasova, 
Fl. Az. Ross, in 7, 1. 

Perennial; rhizome long, slender, violet with brown scales; stem 8—10 cm 
high, leafless or with 1 leaf, erect, covered — like pedicels and petioles — 
with dense glandular hairs, glabrous at base, developing creeping, rooting, 
pubescent shoots; radical leaves 2—6, with petioles of varying length, 
reniform-cordate, 10 — 30 mm long, 15—45 mm broad, covered on both sides 
with sparse white bristly hairs, unequally dentate; flowers in a simple, 
terminal, longish raceme; pedicels 2 — 7 mm long, ebractate; calyx 
patelliform, stellate, 5 mm in diameter, greenish brown, with 5 oblong or 
ovate acuminate sepals, glandular on both sides; petals 4—5 mm long, twice 
as long as sepals, reddish brown, pinnatisect into 8 or 9 filiform lobes; 
stamens 10, with short filaments; pistils with divergent styles, shorter 
than calyx; ovary sunken, yellowish; capsule 2.5 mm long, 3.5 mm broad, 
subglobose, minutely glandular; seeds ovoid, black-brown, shining, 1 mm 
long. June — July. 

Moist, shady pine forests. — W. Siberia: Alt., Irt., Ob; E.Siberia: Yenis., 
Lena-Kol.; Far East : Ze.-Bu., Uss., Uda, Okh. Gen.distr.: Jap.-Ch., 
N.Am. Described from N.America. Type in London. 



Genus 708. CHRYSOSPLENIUH * L. 

L. Sp.pl. (1753) 398. 

Calyx 4-lobed, adnate to ovary at base, greenish or yellow; corolla 
absent; stamens 8, rarely 4, with short filaments inserted on a glandular 
perigynous disk; ovary almost inferior or half -inferior, with short styles; 
capsule unilocular, dehiscing along longitudinal cleft. Perennials or 
annuals with fleshy, glabrous or pubescent stems, with alternate or opposite, 
incised or dentate leaves; radical leaves rosulate or absent, cauline leaves 
few or absent, bracts usually differing from cauline and radical leaves, 
surrounding corymbiform inflorescence, or else flowers solitary, sometimes 
bracts not differing from cauline leaves. 

1. Cauline leaves alternate or absent 2. 

+ Cauline leaves opposite 12. 

2. Stem leafy 3. 

+ Stem leafless, with only radical leaves and bracts present 22. 

3. Flowers in an inflorescence; bracts differing from cauline leaves ... 6. 
+ Flowers solitary, in leaf axils of stem forks 4. 

4. Plants developing creeping shoots; radical leaves not reniform or 

else absent; flowers numerous 5. 

+ Plants developing short, creeping shoots; radical leaves reniform; 

flowers 1 — 3 8. C.komarovii A. Los. 



From the Greek c h r y s o s, gold and s p 1 e n , spleen, referring to the golden flowers and to the use of the 
plant against spleen diseases. 



155 



201 



202 



5. Leaves lobate; radical leaves numerous 9. C. sedakovii Turcz. 

+ Leaves dentate; radical leaves absent ... 11. C. tianschanicum Krassn. 

6. Radical leaves absent; cauline leaves oval, dentate 

10. C. ovalifolium M. B. 

+ Radical leaves numerous; cauline leaves reniform and lobate .... 7. 

7. Plants not giving rise to sterile shoots 8. 

+ Plants with sterile creeping shoots arising in axils of radical 

leaves 10. 

8. Stamens 8 9. 

+ Stamens 4 4. C. tet rand rum (Lund.) F. Fries. 

9. Sepals greenish 3. C. alternifolium L. 

+ Sepals lilac 5. C. beringianum Rose. 

10. Stem 10—25 cm high; shoots longer than stem 

. 6. C. flagelliferum F. Schmidt. 

+ Stem 5 — 9 cm high; shoots shorter than stem 7. C.filipes Kom. 

11. Stem glabrous; leaves deeply dissected into subquadrate, rounded 

lobes l.C. nudicaule Bge. 

+ Stem pubescent with long rufous hairs in lower part and below the 
inflorescence; leaves corymbiform, shallowly incised into 5 — 7 
lobes 3. C. peltatum Turcz. 

12. Plants quite glabrous 13. 

+ Plants more or less pubescent 17. 

13. Stems erect, with creeping sterile shoots arising in axils of lower 

and radical leaves, or rooting shoots absent 14. 

+ Stems prostrate, branching, rooting at the nodes, with erect flowering 

shoots 16. 

14. Flowering stem with one pair of leaves in lower part or leafless; 
leaves inconspicuously dentate or subentire; sterile shoots not 
rooting 15. C. kamtschaticum. 

+ Flowering stem leafy; leaves dentate 15. 

15. Sterile shoots shorter than stem, their terminal leaves not differing 
from all the others; seeds glabrous, smooth .... 14. C. sinense Maxim. 

+ Sterile shoots longer than stem; terminal leaves much larger than 

all the others; seeds covered with small papillae 

16. C. trachyspermum Maxim. 

16. Flowers cyathiform, erect, long-pediceled 17. C.rimosum Kom. 

+ Flowers patelliform, short -pediceled 18. C.dubium J. Gay. 

17. Flowers flat; disk red. Puberulent plants; stem branching in lower 
part 12. C. ramosum Maxim. 

+ Flowers campanulate -infundibular or campanulate -cyathiform. 

Densely pubescent plants 18. 

18. Entire plant pubescent with white hairs, only the bracts subglabrous, 
green 13. C.pilosum Maxim. 

+ Plant pubescent with rufous hairs; flowering stems glabrous; bracts 

glabrous, yellowish green 19. C. baicalense Maxim. 

1. C. nudicaule Bge. in Ldb., Fl. Alt. II (1830) 114; Ldb., PI. Ross. 11,226; 
Kryl., Fl. Zap. Sib. VI, 1432.- Ic. : Ldb., Ic.pl. Fl. Ross. tab. 405; Nekrasova 
in Fl. Az. Ross., No. 7 (1915) 2. 



156 



203 



Perennial; rhizome creeping, thick, with numerous root fibers; stem 
leafless, succulent, glabrous, erect, sulcate, 5—25 cm high; radical leaves 
long-petioled, about as long as stem, rounded -reniform, paler below, 
glabrous or with sparse bristles, incised to l / 4 — % into subquadrate, 
rounded lobes, sometimes broadening in upper part, the blade 1 .5— 5 cm 
broad, 1—3 cm long; bracts smaller, short -petioled, flat or slightly truncate 
at base, with fewer lobes; flowers 5 — 15, infundibular, crowded into a 
corymb, subsessile, 6 — 7 mm in diameter; sepals green, incised to the 
middle into broad, quite truncate, erect lobes, twice as long as broad; 
stamens 8, slightly shorter than calyx; disk yellowish, 8-lobed; capsule 
widely dehiscing by obtuse, rufous valves; seeds 1 mm long, ovoid, black, 
glabrous, shining. Fl. May — June; fr. June — July. (Plate XII, Figure 2). 

Shady banks of mountain rivers and streams, in the alpine zone, 
sometimes occurring below the timberline. — W.Siberia: Alt.; E.Siberia: 
Ang. -Say., Yenis.; Centr.Asia: T. Sh., Dzu. -Tarb. Gen. distr. : Mong., 
China. Described from the Altai. Type in Leningrad. 

2. C.peltatum Turcz., Catal. PI. No. 503 (l 837), Ej., Fl. baic. -dah. I, 
464; Ldb., Fl. Ross.II, 226. — Ic: Franchet, Monogr. in Nouv. Arch. Mus. 
Ser. II (1890) pi. IV A; Nekrasova in Fl. Az. Ross., No. 7 (1915), Fig. 5. 

Perennial; rhizome creeping, with sparse root fibers; stem 5—10 cm 
high, erect, covered in lower part and below inflorescence with sparse 
long rufous hairs; leaves all radical or bracts; petioles of radical leaves 
broadening toward base, covered with long rufous hairs, inserted to 
center of blade, 1— 3 cm long; blade somewhat fleshy, peltate -reniform, 
glabrous above, sparsely hairy below, lighter, 4—8 cm long, 5 — 1 cm 
broad, shallowly incised into 5 — 7 rounded subobtuse lobes; inflorescence 
condensed, corymbiform, few-flowered; bracts rounded -ovate, smaller 
than radical leaves, cuneate, 3— 5 -lobed, the lobes slightly acuminate or 
subobtuse, shallowly incised; pedicels 1 mm long; flowers infundibular, 
open; sepals greenish yellow, broadly ovate, rounded above or slightly 
obtuse, 1.5 mm long, 1 .5— 2.5 mm broad; disk pale green, convex, with very 
prominent lobes; stamens 8, slightly shorter than sepals; ovary with 
short divergent styles; capsule shorter and almost completely adnate 
to calyx, slightly emarginate; seeds glabrous, smooth, shining, 0.75 mm 
long. Fl. May — June, fr. July. 

Rock streams, river and stream banks, in alpine zone. — E. Siberia: 
Ang. -Say. (Tunkinskie Belki [snow-capped mountains]). Gen. distr. : 
Mong. Described from the Tunkinskie Belki, Dzhokoi River, Nukhu-Daban 
Pass. Type in Leningrad. 

3. C.alternifolium L., Sp.pl. ed. I (1753) 398; Ldb., FL Alt. II (1830) 
115; Fl. Ross.II, 226; Shmal'g., Fl. I, 356; Kryl., Fl. Zap. Sib. VI, 1433; 
Kom., Fl. Kamch. II, 220; Kom. and Alis., Opred. rast. Dal'nevost. kraya, I 
(1931) 609.- C. nivale Schur, Enum. PL Transsilv. (1866) 241.- Ic. : 
Nekrasova in Fl. Az. Ross., No. 7 (1915), Fig. 6; Rchb., Ic. Fl. Germ, et Helv. 
XXVI (1898) 129.- Exs.: PL Finl. Exs., No. 259. 

Perennial; rhizome slender, pale brown, with numerous root fibers; stem 
solitary, less often several, erect, glabrous or with sparse white or rufous 
hairs in lower part, 5—15 cm high, with 1—3 alternate leaves; leaves 



157 



204 



somewhat fleshy, light green, paler below, rounded -reniform, sometimes 
subquadrate, with deeply cordate base, shallowly incised on the margin 
into rounded or subquadrate lobes, covered on both sides with sparse 
hairs or glabrous; radical leaves few, 7—25 mm long, 10— 35 mm broad, with 
petioles to 5 cm long; cauline leaves smaller, short -petioled, with flatter 
base or even cuneate; bracts 4— 15 mm in diameter, yellowish, with fewer 
lobes, sometimes entire, broadly cuneate, with broad, glabrous petioles 
1—2 mm long; inflorescence flat-corymbiform; flowers numerous, with 
petioles 1 mm long, elongating in fruit; sepals ovate, flat lobes, golden- 
yellow inside, 1.5—2 mm long; disk fleshy; stamens 8, shorter than 
calyx; styles short; capsule as long as calyx, with broad and rather small 
notch; seeds oblong -ovoid, glabrous, smooth and lustrous. Fl. April- 
June, fr. May —July. 

Moist and shady sites, among shrubs in flood meadows, on banks of 
rivers, streams, and lakes in the Arctic and forest zones; Arctic: Nov. Z., 
Arc. Eur., Arc. Sib. ; European part : Kar. : Lap., Dv. -Pech., Lad. -Ilm., U. V., 
V. -Kama, V. -Don, M. Dnp., L. Don (N. part); Caucasus: Cisc, E. and W. 
Transc; W.Siberia: Ob, U. Tob., Irt., Alt.; E.Siberia: Yenis., Ang. -Say., 
Lena-Kol., Dau.; Far East: Kamch., Okh., Uda, Uss., Ze. -Bu. Gen. distr. : 
Arc, Centr. Eur., Scand., Jap. -Ch., Mong., N. Am., Bal. -As. Min. Described 
from Switzerland. Type in London. 

4. C.tetrandrum (Lund) Th. Fries, Bot. Not., Ko. 12 (1858) 193.- 
C.alternifolium var. tetrandrum Lund ex Malmgren in Bot. Not., 
No. 3 (1846)39,78; Kryl., Fl. Zap. Sib. VI, 1433. 

Perennial, glabrous, sometimes the young radical leaves short -hairy; 
stem erect, 5 — 15 cm high, succulent, leafy; radical leaf petioles 2—3 times 
as long as blade, rounded -reniform, with cordate base, broader than long, 
10—15 mm broad, 6—12 mm long, with 5—9 rounded -oval or subobtuse lobes; 
cauline leaves smaller, 3—5 -lobed, with shorter petioles; bracts trilobate, 
cuneate; inflorescence condensed, corymbiform, few -flowered; pedicels 
1— 1.5 mm long; calyx greenish yellow, with oval-triangular sepals; 
stamens 4; disk greenish, 4 -lobed; styles erect, short; capsule as long as 
calyx, with broadly oval obtuse lobes; seeds shining, smooth, glabrous, 
light brown. June — July. 

Stream and river banks in the tundra. — Arctic : Arc. Eur., Nov. Z., Arc. 
Sib. Gen. distr. : Ber.,N.Am. Described from Scandinavia. 

5. C.beringianum Rose in Bot.Gaz.23 (1897) 275; Hulten, Fl. oj Kamtch. 
Ill (1929) 32.- Ic: Macoun, PI. Pribil. Isl. Ill (1899) tab. 9. 

Perennial; rhizome 2.5 — 5cm long, creeping, giving rise to many long, 
fibrous roots; stems few, 2.5—5 cm high, glabrous, leafless, less often 
bearing 1 leaf each; radical leaves in a dense rosette; blade pale and 
glabrous below, dark green and slightly hairy above, reniform, 6—11 mm 
long and as broad, with 4 or 5 glandiferous teeth; petioles slender, slightly 
broadening downward, 1.5—4.5 cm long, with long violet hairs on the margin 
(especially in lower part); bracts entire or trilobate; flowers violet, 
sometimes green at beginning of anthesis; sepals suborbicular; disk 
very fleshy, deeply incised into 8 lobes; fruiting calyx growing and 
becoming tubular; capsule bilobate; seeds oblong, 0.5 mm long, shining, 
finely reticulate. July. 



158 



Volcano slopes.— Far East: Kamch. (Klyuchevskaya Sopka). Gen. distr.: 
Ber., Alaska. Described from the shores of Saint Paul Harbor. Type in 
New York. 

*C.wrightii Franch. et Sav., Enum. PI. Jap. II (1879) 356 in adnot.; Kom., 
Fl. Kamch. II (1929) 221.- Ic: Franch. Monogr. in Nouv. Arch. Mus. 3-e 
ser. PL V, 13; Kekrasova in Fl. Az. Ross., No. 7 (l 915), Fig. 7. 

Perennial; stem erect, covered with rufous hairs; leaves succulent, 
3—7 cm in diameter, orbicular, dentate, with cuneate base, rufous -hairy 
on the margin; radical leaves with long petioles, the bracts short - 
petioled, all petioles pubescent; inflorescence few-flowered; flowers 
subsessile; calyx green, with orbicular sepals; disk thin, with 8 obscure 
lobes; stamens 8; capsule as long as calyx; seeds glabrous, narrow, 
smooth. 

Note. Collected by Wright, a member of the Reinhold and Rodgers 
expedition, probably on the west coast of Kamchatka; there is no precise 
data as to where the specimen was collected; this species has been 
included in the Flora of Kamchatka on the basis of that single specimen; 
the type is preserved in the Paris Museum. V. L. Komarov is doubtful as 
to the occurrence of this specimen in Kamchatka, for nobody else has found 
it there. Hulten (Fl. Kamch. Ill (1929) 35) puts forward the hypothesis 
that this species was collected in Sakhalin, Hokkaido, or the Kurile Islands, 
and excludes it from the Flora of Kamchatka. On account of these indefinite 
views, it seems risky to include C.wrightii Franch. et Sav. in the Flora 
of the USSR. 

6. C. flagelliferum F. Schmidt, Reise in Amurl. und Sachal. (1868) 134; 
Tr. Sib. Eksped.(l879) 146; Kom., Fl. Manch. II (1904) 426; Kom. and Alis., 
Opred. rast. Dal'nevost., kraya I (l 931 ) 609. — Ic. : Franch. Monogr. in Nouv. 
Arch. Mus. 3 Ser. (1890) II, PI. Ill; Nekrasova in Fl. Az. Ross. No. 7 (1915), 
Fig. 10. 

Perennial, with creeping, branching, leafy shoots, with terminal rosettes, 
rooting at the nodes; flowering stems 2 — 3, to 20 cm high, tetragonal, reddish, 
sparsely pubescent with reddish hairs, pseudo-dichasially branching in 
upper part; radical and cauline leaves thin, paler below, rounded -reniform, 
glabrous or minutely hairy, with obtuse, rounded, glandiferous teeth on the 
margin; radical leaves ca. 5 cm long, 6 cm broad, the petioles longer than 
the blade, pubescent with rufous or whitish hairs, broadening downward, 
ciliate; cauline leaves alternate, 1—2 cm in diameter, palmatisect, entire 
at base or slightly cuneate, short -petioled; bracts smaller than cauline 
leaves, oblong, somewhat oblique, 3—5 -lobed, glabrous, cuneately tapering 
to short glabrous petiole; flowers few, open, flat, subsessile in stem forks 
and in axils of upper leaves, forming an inflorescence composed of 2 
peduncles; sepals dissimilar, the largest rounded -rhomboid or oval- 
rhomboid with recurved margins, 1.5 mm long, 1 mm broad, all sepals 
yellowish; stamens 8, with minute filaments; disk greenish, obscurely 
8 -lobed, fleshy; ovary sunken, with 2 erect styles slightly longer than 
stamens; capsule widely dehiscing by oval lobes; seeds shining, smooth, 
sometimes covered with slight pubescence. Fl. May, fr. June. (Plate XII, 
Figure 4). 



159 



207 



Mixed moist forests, on moist soils, often along streamlet banks. — 
Far East: Ze. -Bu., Uss., Sakh. Gen. distr. : Mong., Jap. -Ch. Described 
from the Bureya Range. Type in Leningrad. 

7. C.filipes Kom. in Fedde Repert.sp. n. IX (1911) 393; Kryl., Fl. 
Zap. Sib. VI, 1435.- Ic: Nekrasova in Fl. Az. Ross., No. 7 (1935), Fig. 9. 

Perennial, glabrous, delicate; rhizome slender, long; stem 2 — 9 cm 
high, slender, weak, not branching, leafless or with 2—3 small leaves, 
giving rise to sterile creeping shoots below; radical leaves thin, the 
blade longer than or as long as petiole, reniform or orbicular, slightly 
tapering toward base, with 4 or 5 rounded glanduliferous teeth on the 
margin, 4—8 mm long, 6—10 mm broad, paler below; cauline leaves alternate, 
very small, cuneate; bracts larger than cauline leaves, short -petioled; 
inflorescence 3— 5 -flowered, the pedicels 0.5—1 mm long; sepals greenish, 
orbicular or rounded -ovate, 0.8—1 mm long, 1—1.3 mm broad; stamens 8, 
with short filaments; disk fleshy, green; ovary not sunken, with divergent 
styles; seeds brown, dull, covered with minute villi. Fl. May, fr. June. 

Mountain forests, shady, moist stream banks, rock crevices. — 
W.Siberia: Alt.; E.Siberia: Ang.-Say. Endemic. Described from the 
Kulumys Range. Type in Leningrad. 

8. C. komarovii A. Los. sp. nova in Addenda VIII, p. 374. 
Perennial; stem 2—5 cm high, slender, glabrous, giving rise to few 

slender, short, curved shoots toward end of anthesis, with a terminal 
rooting rosette; radical leaves rosulate, reniform, coarsely crenate- 
dentate, with glanduliferous teeth, glabrous below, glaucescent, covered 
with minute bristly hairs above, somewhat fleshy, 0.5—1.5 cm long, 
shorter than broad; cauline leaves 1 or 2, alternate, smaller than the 
radical, with cuneate base and smaller teeth; bracts glabrous, smaller 
than cauline leaves, cuneate, trilobate; flowers solitary, with 1—1.5 mm 
long pedicels, 1 or 2 on stem in axils of bracts, flat; sepals greenish, 
oval -triangular; disk very slightly protruding, not fleshy, dull, 1 mm 
long; stamens with yellow anthers and minute filaments; ovary sunken; 
styles erect. July. 

Among stones, along mountain streams; in forests.— Far East: Uss. 
(Suputinka). Endemic. Described from the Suputinka River. Type in 
Leningrad. 

9. C.sedakowii Turcz., Fl. baic.-dah. I (1842) 464; Bull. Soc. Nat. Mosc. 
XVII,273; Kryl., FL Zap. Sib. VI, 1435.- rupifraga sedakowii Turcz. 
in sched. — Ic. : Franch., Monogr. in Nouv. Arch. Mus. 3-e ser. Ill (1890) 
PL VIA.et Al; Nekrasova in FL Az. Ross., No. 7 (l 915), Fig. 8. 

Perennial, slender, very delicate, quite glabrous; roots long, slender; 
stem bearing at base a small reserve, wintering bud (bulbil) surrounded by 
broadened petiole bases, filiform, branching, 5—15 cm high, leafy; all 
leaves similar, petiolate, with prominent veins, nearly twice as broad as 
long, with slightly cuneate or slightly obtuse base, with 3—5 oval lobes; 
cauline leaves alternate; petioles of various length, those of radical leaves 
with fimbriate hairy stipules; flowers few, solitary, with pedicels to 3 cm 
long, nodding in fruit, in stem forks or in axils of upper cauline leaves; 



160 



sepals dingy white or greenish, rounded -oval, 1 mm in diameter, erect; 
disk thin, pale green; stamens 4, rarely 8, shorter than sepals; ovary 
orbicular, half-sunken; styles erect, shorter than stamens; capsule 
slightly emarginate; seeds smooth, glabrous, ovoid, 4 mm long. Fl. May, 
fr. June- July. (Plate XII, Figure 1 ). 

Moist shady sites (under rocks, boulders, etc.), taiga dominated by 
Abies s ibi ri c a, and sparse deciduous forests. — W. Siberia: Alt. 
(Lake Teletskoe); E.Siberia: Ang.-Say., Dau. Gen. distr. : N. Mong. 
Described from the vicinity of Nerchinsk. Type in Leningrad. 

10. C.ovalifolium M. B.ex Bunge in Ldb., Fl. Alt. II (1830) 115; Ldb., 
Fl. Ross. II, 227; Kryl., Fl. Zap. Sib. VI, 1434. - Ic. : Ldb., Ic. pi. Fl. Ross., 
tab. 474 (1834); Franch., Monogr. in Nouv. Arch. Mus. 3-e ser. Ill (l 890) 
PI. VI B; Nekrasova, in Fl. Az. Ross., No. 7 (1915), Fig. 4. 

Perennial; rhizome creeping, rufous, with slender roots and with 
brown scales above; flowering stem erect, curved or straight, 10—16 cm 
high, finely sulcate, leafy; sterile shoots more densely leafy, straight, 
20— 25 cm high; radical leaves absent; cauline leaves alternate, green, 
paler below, glabrous, elliptic, rounded -obovate or ovate, broadly cuneately 
tapering to petiole, shorter than the blade, serrate -dentate, the teeth with 
forward -bending tips, obtuse; leaves on shoots 1— 4 cm long, 0.5— 2.5 cm 
broad, those on flowering stem smaller; bracts similar to cauline leaves 
but with fewer teeth; flowers in a branching corymb, with 0.5— 4 mm pedicels; 
sepals green, broadly ovate, obtuse, 1.5 mm long; disk pale, fleshy; 
stamens 8, shorter than sepals; capsule with acute divergent lobes; seeds 
subglobose, 0.5 mm in diameter, smooth, dull with short pubescence. 
Fl. May -June, fr. July. 

Spruce, Siberian stone pine, and fir forests (dominated by Abies 
sibir ica), not penetrating into mountains.— W.Siberia: Alt.; E.Siberia: 
Yenis. Endemic. Described from the Altai. Type in Leningrad. 

11. C. thianschanicum Krassn., Opyt razvitiya Fl. Zap. Tyan' -Shanya, 
Zap. R. G. O-va.XIX (l 888) 132 and 137, Plate I; Descr. PI. nov. in Scr. 
Bot. II (1889) 16. 

Perennial; roots thick, cordlike; stems bearing at base a large oval 
reserve winter bud, 6—20 cm high, branching from the middle, leafy, the 
leaves replaced in lower part by brown scales; radical leaves absent; 
cauline leaves alternate, similar, pale green, paler below, thin, ovate or 
broadly ovate, with cuneate base, serrate -dentate, 0.5—1.5 cm long and broad; 
cauline leaves long-petioled; bracts and upper cauline leaves short -petioled; 
flowers solitary, in stem forks and in axils of upper leaves, erect or nodding, 
with slender pedicels to 20 mm long; sepals greenish, triangular -ovate, 2 mm 
long, 2.5— 3 mm broad, 3 -nerved, with acuminate lobes forming rounded notch 
between adjacent sepals; stamens 8; disk fleshy, 8 -lobed, yellowish; capsule 
not longer than calyx, with short lobes; seeds oblong-ovoid, glabrous, smooth, 
shining. Fl. July, fr. August. (Plate XII, Figure 3). 

Moist rocks above the timberline. — Centr. Asia: T. Sh. Gen. distr. : 
Dzu.-Kash. (Kuldja). Described from the Ak-Su River. Type in Leningrad. 

12. C.ramosum Maxim., Prim. Fl. Amur. (1859) 121; Kom., Fl. Manch. H 
(1904) 42; Kom. and Alis., Opr. rast. Dal'nevost. kraya I (1931 ) 610.- 



161 



C. opp o s it i f o 1 i um Trautv. und Meyer., Fl. ochot. (l 856) 42 (non Cham, 
et Schlecht.). — C. y e z o e n s e Franch. et Sav. Enum. PL Jap. II (1879) 
355 et 649.- Ic: Franch., Monogr. in Nouv. Arch. Mus. 3-e Ser. II (1890) 
Pl.IVB. 

Perennial; rhizome slender, creeping; stem opposite -branching from the 
base or the middle, 6—20 cm high, slightly angular, pubescent with sparse 
remote hairs, separating in upper part into 2 peduncles; lateral shoots 
sterile, often longer than the flowering shoot, ascending or prostrate, rooting 
at lower nodes; leaves opposite, their amplexicaul petioles pubescent 
at base, the blade broadly rounded, blunted, slightly cuneate toward base, 
7— 8 mm long, 12—13 mm broad, serrate-dentate, with glandiferous teeth; 
-, q sterile shoots developing terminal rosettes of larger leaves; bracts smaller, 
always glabrous, with slightly attenuate apex; inflorescence with subglabrous 
branches; flowers in a corymb, 3—7 per peduncle, subsessile or sessile; 
inflorescence quite flat; sepals green, broadly ovate -cordate, the 2 opposite 
larger, hence the flowers somewhat elongated and oval in outline; large 
sepals 1 .5—2.5 mm long, 1 .5—2 mm broad, small sepals 1—2 mm long, 1 mm 
broad; stamens 8, with minute filaments; anthers orange; disk red, fleshy, 
strongly convex, with 8 deep lobes; ovary flattened, half-sunken, with short 
erect styles; capsule with ovate divergent lobes, widely dehiscing; seeds 
oblong, glabrous, smooth, 0.5—1 mm long. FL May, fr. June. (Plate XII, 
Figure 5). 

Forests, river and stream banks.— Far East: Ze. -Bu., Uss., Uda. 
Gen. distr.: Jap. -Ch. (japan, Korea, Manchuria). Described from 
De-Kastri Bay, near the village of Ngalmar. Type in Leningrad. 

13. C.pilosum Maxim., Prim., Fl. Amur. (1859) 122; Kom., Fl. Manch. II, 
424 ; Kom. and Alis., Opr. rast. Dal'nevost. kraya I, 609. — Ic. : Franch. in 
Nouv. Arch. Mus. 3-e Se'r. Ill (1890) PL HI D.-C. 

Perennial; stems to 20 cm high, numerous, branching from the middle, 
tetragonal, densely covered with whitish, later rufous, horizontally spreading 
hairs; in fall the plant developing sterile creeping shoots with large 
rooting rosettes; cauline leaves opposite, suborbicular, cuneate at base, 
crenate, 0.5—1 .5 cm in diameter, covered on both sides with scattered short 
hairs, lighter below with transparent veins, the densely pubescent petioles 
as long as the blade; leaves on sterile shoots to 4 cm in diameter; bracts 
glabrous, covered with sparse hairs on the margin, oval -rhomboid, oval or 
oval -triangular, irregularly serrate -dentate, the smallest bracts entire, the 
short petioles pubescent below and onthe margin; inflorescence consisting 
of 2 branching peduncles each bearing up to 12 flowers, the largest first 
flowers — always greener than the upper ones — usually in inflorescence 
forks; pedicels of varying length; flowers patelliform; calyx connate to 
the middle, the sepals semiorbicular, concave and reflexed -tipped, yellow - 
green; stamens 8, shorter than calyx, the filaments slightly dilated toward 
base; disk greenish, not convex, with 8 rounded lobes; ovary sunken, 
rounded -oval, with erect style; capsule with dissimilar, arched -upcurved 
acuminate lobes, twice as large as calyx; seeds ovoid, ribbed, tuberculate 
along the ribs. Fl. May, fr. June. 

Mixed forests, mountain valleys, riverbanks, mossy thickets, shore 
sandS.— Far East: Uss. Gen. distr. : Jap. -Ch. (Manchuria, Korea, China). 
210 Described from cliffs opposite the village of Tsyanka on the lower Amur. 
Type in Leningrad. 



162 



211 




PLATE XII. 1-Chrysosplenium sedakowii Turcz., a) flower from the side, b) flower from above; 
2 — C.nud icaule Bge., a) flower; 3 - C. th i a ns cha ni c um Krassn., a) flower from above; 4 — 
C. flagelliferu m F.Schmidt, a) flower from the side, b) flower from above, c) leaf teeth and glands, 
d) seed; 5 — C.ramosum Maxim., a) flower from the side, b) flower from above, c) seed. 



163 



:i3 



14. C. sinicum Maxim., Diagn. PL nov. asiat. in Mel. Biol. IX (1877) 769; 
Kom., Fl. Manch. II (1904) 424; Kom. and Alis., Opred. rast. Dal'nevost. 
kraya I (l 931 ) 609. — Ic. : Franch., Monogr. in Nouv. Arch. 3 -e ser. Ill 
(1890) PI. VI E. 

Perennial; rhizome short, fibrous; stem erect, glabrous, 3—12 cm high, 
in lower part developing sterile shoots arising from axils of lower and 
radical leaves; leaves opposite, slender, glabrous, pale below, all leaves 
denticulate -serrulate, the radical 2—3 cm long, 6 — 15 mm broad, oblong- 
ovate, with cuneate base, short -petioled; cauline leaves 2 or 3 pairs in 
terminal rosettes at tips of sterile shoots, orbicular, broadly cuneate or 
slightly cordate, their petioles as long as or longer than the blade; bracts 
yellowish, broadly elliptic, 7—12 mm long, 4—7 mm broad, short -petioled; 
inflorescence composed of 2 short branches forming a globose corymb; 
flowers few, subsessile; calyx campanulate, with orbicular subobtuse, 
yellow sepals, one pair 2 mm in diameter, the other opposite pair 
1.5—2 mm long, 1—1.5 mm broad; stamens 8, as long as or shorter 
than calyx; ovary sunken, with erect divergent styles; disk pale, not 
fleshy, 8-lobed; capsule not exceeding calyx, with broadly ovate, acuminate, 
lobes; seeds glabrous, smooth, ovoid, 0.75 — 1 mm long. May. 

Forests.— Far East: Ze.-Bu. (Bureya Range), Uss. (Barabash natural 
boundary area). Gen. distr. : Jap. -Ch. (Manchuria, Kansu Province). 
Described from Kansu Province. Type in Leningrad. 

15. C. kamtschaticum Fisch. in DC, Prodr. IV (1830) 48; Ldb., Fl. Ross. 
11,227; Kom., Fl.Kamch. H (1929) 221.- C.oppositifolium Cham, et 
Schlecht., Linnaea VI (1831)557.— Ic: Franch. , Monogr. in Nouv. Arch. 
Mus. 3-e ser., Ill (1890) PL lie; Nekrasova in Fl. Az. Ross., No. 7 (l 915), 
Fig. 14. 

Perennial; rhizome short; stem erect or slightly flexuous, leafless or 
with 1 or 2 pairs of leaves, glabrous, branching in the inflorescence, 4—20 cm 
high, developing short nonrooting sterile shoots with terminal rosette, 
arising from axils of radical or cauline leaves; leaves firm, darker above, 
glabrous, the radical rounded or broadly oboval, broadly cuneate, 7—15 mm 
in diameter, obscurely dentate or entire-margined, with glabrous petioles 
3 mm long; cauline leaves opposite, flabellate, 7—8 mm long, 5 — 7 mm broad, 
dentate in upper part, with petioles to 7 mm long; rosette leaves of 
sterile shoots orbicular, broadly cuneate or cordate, 5— 15 mm in diameter, 
short -petioled; bracts broadly obovate, crenate-dentate, 9— 18 mm long, 
7—15 mm broad, with petioles to 5 mm long; inflorescence condensed, 
corymb iform, few -flowered; pedicels 1.5—5 mm long; flowers greenish; 
sepals broadly oval, 1.5 mm; disk 8-lobed; stamens 9, shorter than calyx; 
styles divergent; capsule deeply dissected, with unequal,' arched -upcurved 
lobes 2—3 times as long as calyx; seeds ovoid, narrowed toward the ends, 
with a deep groove and 11—13 obtuse longitudinal ribs connected by slender, 
approximate, transverse lines. June. 

In shade, in stony beds of mountain -spring brooks, spring bogs with 
stony bottoms, in groups and thickets.— Arctic: Arc. Sib., Chuk., An.; 
Far East: Kamch., Sakh. Gen. distr. : Japan, Kurile Islands. Described 
from Kamchatka. Type in Geneva. 



164 



16. C. trachyspermum Maxim., Diagn. PL nov. asiat. in Mel. Biol. XI 
(1881)226; Kom., Fl. Manch. IV (1904) 422. - Ic. : Franch., Monogr. in 
Nouv. Arch. Mus. 3-e se'r. Ill (1890) PI. I B; Nekrasova in Fl. Az. Ross., No. 7 
(1915), Fig. 12. 

Perennial; rhizome slender, short; stem erect, glabrous, 7—15 cm 
high, leafy, developing in lower part ascending, leafy, sterile shoots rooting 
at the nodes, with a terminal rosette of large leaves, the shoots as long as 
or longer than stem; cauline leaves and bracts slender, delicate, glabrous, 
with delicate, dark, branching veins; cauline leaves opposite, 2—4 pairs, 
rounded -ovate, 6—10 mm long, 6—9 mm broad, slightly obtuse, serrate - 
dentate, broadly cuneate, tapering to a glabrous petiole 3—12 mm long; 
leaves at tips of shoots ovate, rounded -ovate, or oblong-ovate, cuneate, 
serrate -dentate, entire in lower part, with rufous ciliate hairs at base, 
the petioles 5— 10 mm long in fall specimens, the leaves often firm; bracts 
oblong-ovate, 10—15 mm long, 3—7 mm broad, dentate, tapering upward, 
with short glabrous petioles; inflorescence a regular, gJobose corymb 
with 2 or 3 short peduncles; flowers few, subsessile; pedicels elongating 
in fruit to 5 mm; calyx greenish, broadly campanulate, with orbicular or 
rounded -quadrate concave lobes, 2 mm long, 1.5 mm broad; disk thin, pale; 
stamens 8, shorter than calyx; styles divergent and protruding from calyx; 
capsule with unequal, arched -upcurved, acuminate lobes twice as long 
as calyx; seeds dark, smooth, shining, covered with small papillae, acuminate, 
0.5—0.7 mm long. Fl. May, fr. June— July. 

Mixed shady moist forests; often forming thickets.— Far East: Ze.-Bu., 
Uss. Gen. distr. : Jap.-Ch. Described from the Bureya Mountains. Type 
in Leningrad. 

17. C.rimosum Kom. in Fedde, Repert. sp. nov. XIII (1914) 168; Kom., 
Fl.Kamch. II (1929) 222. 

Perennial; rhizome creeping; stem ascending at base, erect higher 
up, 5—10 cm high, glabrous, with prostrate shoots arising from axils of 
lower leaves; sterile shoots rooting at the nodes, opposite -branching; 
radical leaves rosulate, semiorbicular or flabellate, short -petioled, slightly 
crenate; cauline leaves few, opposite, slightly cordate, subsessile, with 
deeper teeth; leaves on sterile shoots numerous, the petioles half as 
long as blade; all leaves glabrous, fleshy, light green, the blade 0.3—1 cm 
in diameter; bracts rounded -oval to oval, with fewer teeth, often obscurely 
trilobate; inflorescence condensed, composed of 3—5 erect flowers; 
pedicels 0.2—0.6 cm long; calyx campanulate, with green, oboval sepals, 
2 mm long; disk green, flat; capsule with erect lobes longer than calyx; 
seeds dull, ovoid, rugose. August. 

Pebbles, gravelly slopes, rocks, along streams in the alpine zone. — 
Far East: Kamch. Endemic. Described from Kamchatka. Type in 
Leningrad. 

18. C.dubium J. Gay in DC., Prodr. IV (1830) 48.-C.macrocarpum 
Cham, et Schlecht. in Linnaea VI (1831) 558; Boiss., FL Or. II (1872) 813; 
Grossg., Fl.Kavk. II (1930) 24.- Ic. : Franch., Monogr. in Nouv. Arch. Mus. 
3-e ser. Ill (1890) PL II D. - Exs. : Fl. cauc. exs., No. 93. 

Perennial, completely glabrous; rhizome long, creeping; stem decumbent 
or ascending, leafy, rooting at the nodes; flowering shoots ascending, 



165 



215 



216 



10—20 cm high, leafy; leaves fleshy, paler below, opposite, petiolate, 
elliptic, ovate, or obovate, rounded above, with rounded obtuse or cuneate 
base, dentate; upper leaves of sterile shoots suborbicular, to 2.5 cm in 
diameter, the lower half as long and broad; bracts 8 mm long, rounded -ovate, 
broadly cuneate; inflorescence branching into 2 corymbs; flowers 
yellowish green, small, to 4 mm in diameter; sepals spreading at anthesis, 
oval, 2 mm long; stamens as long as calyx; disk fleshy, 8-lobed, rufous; 
styles short, erect; capsule with oblong lobes, twice as long as calyx; 
seeds 1 mm long, broadly ovoid, with 15 longitudinal rows of clavate villi 
(papillae and transverse lines between them). April— May. 

Gorges, stream banks, on moist soils, in the shade. — Caucasus: 
W. Transc. Gen.distr.: Arm. -Kurd., Bal. -As. Min., W. Med. (Italy). 
Described from Asia Minor. Type in Geneva. 

19. C.baicalense Maxim., Bull. Soc. Nat. Mosc. (1879), No. 22; Kom., 
Fl. Manch. II (1904) 423.- Ic: Franch., Monogr. in Nouv. Arch. Mus. 3-e 
ser., Ill (1890), Fl. 2 A; Nekrasova, Fl. Az. Ross., No. 7 (1915), Fig. 13. 

Perennial; rhizome long, creeping, covered with brown fibers; stem 
prostrate, sulcate, pubescent with long rufous hairs, with long, sterile, leafy, 
rooting shoots; flowering stem erect, glabrous, usually leafless, to 12 cm 
high, light yellow, arising in forks of creeping shoots from leaf axils; 
leaves opposite, orbicular, 15—30 mm in diameter, covered on both sides 
and on the margin with sparse rufous hairs, coarsely but shallowly crenate, 
with cuneate base, paler below, with petioles shorter than the blade; 
bracts glabrous, light yellow -green, very obscurely crenate, slightly larger 
than shoot leaves; flowers few, in an approximate corymb, campanulate- 
infundibular; sepals greenish, orbicular, opposite, dissimilar, the larger 
2.5 mm long, 3 mm broad, the smaller 2 mm long, 1 .5 mm broad; stamens 
shorter than calyx; disk pale, not convex, 8-lobed; styles divergent; 
capsule with acuminate lobes much longer than calyx; seeds slightly 
shining, sulcate and verruculose, narrowed toward the ends. June — July. 

Rock streams, balds, stream banks. — E.Siberia: Ang.-Say. Endemic. 
Described from the Baikal shores, near the village of Kultuk. Type in 
Leningrad. 



Tribe 2. PARNASSIEAE S. F. Gray, Nat. Arr. Brit. PL II (1821 ) 623 (fam.); 
Engl. u. Prantl, Nat. Pflanzenfam. Ill, 2a (1890) 66.- Carpels 3 or 4, 
connate; styles absent or short. Fruit a 3- or 4 -valved capsule with parietal 
placentation. 

Genus 709. PARNASSIA * L. 

L.Gen.Pl.ed.V (1754) 133. 

Perennial glabrous herbaceous plants with short rhizome, fibrous roots, 
simple numerous or solitary stems, entire petiolate radical leaves and 
sessile cauline leaves. Flowers white, solitary at the summit; calyx 
of 5 free or fused lobes; petals 5; stamens 5, alternating with 5 stalked 
palmate appendage -staminodes, divided into lobes and often glandiferous; 
pistils with pyramidal or oval ovary and 3 or 4 sessile stigmas; capsule 
unilocular, 3 - or 4— valved; seeds numerous, attached to walls. 

• This name was given due to the occurrence of this plant on Mt. Parnassus. 

166 



1. Stem with 1 or 2 leaves; petals longer than calyx 2. 

+ Stems leafless; petals as long as or shorter than calyx 

2. P. kotzebuei Cham, et Schlecht. 

2. Calyx with free sepals; staminodes with numerous (6—23) filiform 
glandiferous lobes 1 . P. palustris L. 

+ Calyx connate at base or to the middle; staminodes bi- or 

tripartite 3. 

3. Cauline leaves 2; staminodes bi-partite, eglandulose; calyx connate 

at base 3. P. bifolia Nekras. 

+ Cauline leaf solitary; staminodes tripartite; middle lobule with a 

gland; calyx connate to the middle . 4. P. laxmanni Pall. 

1. P. palustris L., Sp.pl. ed. I (1753) 273; Ldb., Fl. Ross. I, 262; 
Nekrasova, Fl. Az. Ross., No. 11 (1917) 20; Kom., FL Kamch. II (1929) 2 3; 
Kryl., Fl. Zap. Sib. VI, 1436.- P. ciliata Gilib., Fl. Lith. V (1782) 139.- 
Ic. : Nekrasova, 1. c, Plate III B (var. u s s u r i e n s i s Kom.), Plate IX A 
(var. tenuis Wahlb.). 

Perennial, completely glabrous; stems solitary or few (to 15), erect, 
not branching, faintly ribbed, 8—40 cm high; radical leaves oval with 
cordate base, subobtuse, entire, with petioles to 7 cm long, the blade to 
4 cm long, 3 cm broad; cauline leaf 1, of same size and shape, sessile, 
somewhat amplexicaul; flower solitary at the summit, 1.5—4 cm in 
diameter; calyx dissected to base; sepals oval-triangular -lanceolate or 
lanceolate -linear, bluntly acuminate, always shorter than corolla; petals 
5—15, white, with longitudinal yellowish brown or greenish nerves, ovate, 
broadly -ovate or elliptic, obtuse or somewhat acuminate, cuneately tapering 
toward base, 5—15 mm long, 4—12 mm broad; staminodes stalked, broadly - 
oval, with 6—23 filiform glandiferous lobes; stamens equal, with thick 
filaments tapering upward and with white bilocular anthers as long as 
filaments; ovary ovate -pyramidal or ovate -rounded, white or reddish 
with violet dots, with 4 sessile stigmas; capsule dehiscing by 4 valves; 
seeds light brown, numerous, ellipsoid, 0.5 mm long, with lighter border. 
July — August. 

Damp and boggy soils, banks of small river and streams.— European 
part: all regions except Bl., Crim., rarely in the steppe zone; Caucasus: 
all regions except Tal.; Siberia: all regions; Centr.Asia: Balkh., 
Dzu. -Tarb., T. Sh., Pam. -Al. Gen. distr. : Centr. Eur., Atl. Eur., Scand., 
Arc. Eur., Med., Bal. -As. Min., Mong., Jap. -Ch., Tib., N. Am., Ber. 
Described from Europe. Type in London. 

Note. Very variable species; the following characters vary throughout 
the distribution area: overall size of plant, size of leaves, shape of petals, 
number of radical leaves, shape of cauline leaf and its position on stem, 
number of staminode lobes. Different combinations of these characters 
can be observed in various geographical regions, enabling the division 
of the species into races. These races are connected by transitions; we 
therefore prefer to call them forms rather than separate them as 
independent species. 

F.typica Trautv. in A.H.P. V, 1 (1877) 29. — Cauline leaves broadly 
ovate, with cordate, amplexicaul base; sepals oval -triangular, oval, with 
rounded apex and numerous nerves; staminodes 9— 13-lobed; ovary slightly 
tapering into beak. Throughout the distribution area. 

F. tenuis (Wahlb.) Nakras., I.e., 27.- P. tenuis Wahlb., Fl. Lap. (1812) 
74.- P. mult is eta Fernald in Rhodora 28 (1926) 211.- Radical leaves 

167 



few; cauline leaves oval, narrowly oval, and oval -lanceolate, lanceolate in 
extreme forms, always obtuse and with somewhat oblique base; stems 
elongated; sepals lance-linear, few-nerved; petals tapering upward, 

5 — 7 -nerved; staminodes 9— 11-lobed; ovary elongated into beak.— Arctic: 
Arc. Sib. to Kamch; E.Siberia: Lena-Kol., Dau; Far East: Ze.-Bu. 

Gen. distr.: Arc. Eur., N. Am. 

F.ussuriensis Kom. exNekras., 1. c, 28. — Large, robust plants with 
large flowers and coarse, nearly coriaceous leaves; cauline leaves usually 
above the middle; stems solitary; staminodes 17— 21-lobed; ovary 
rounded -oval, not tapering into beak.— Far East: Uss., Ze.-Bu., Uda. 

F.caucasica A.Los., f. nov. — Cauline leaves broadly ovate, acuminate, 
cordate and amplexicaul at base, with 5 very prominent nerves; petals 
oval, with pale nerves; staminodes 5— 7-lobed; ovary rounded, not tapering 
into beak. Caucasus. 

In addition, note should be taken of a Polar -Arctic form, called by 
>„ 1 o V. L. Komarov in his herbarium for Kamchatka f. alpina, — it is the plant 
described by Ruprecht in Beitr. Pflanz. Russ. Reich. II (1845) 23 as 
P. obtusiflora — a low, stocky, shrubby plant with large flowers and 
pale leaves. 

2. P. kotzebuei Cham, et Schlecht. in Linnaea I (1826) 549; Ldb., Fl. 
Ross. I, 264; Kom., Fl. Kamch. II (1929) 224.- P.parvif lor a var. 
kotzebuei Engl., Bot. Jahrb. XIX (1895) 378,473.- Ic. : Britton et Brown, 
Fl. of N. Un. St. II, f. 1854 (1897) 2, f. 2147 (1913). 

Perennial; stem erect, 4—15 cm high, solitary, less often 4, slightly 
ribbed; radical leaves 4 or 5, ovate, with cordate base or broadly cuneate- 
tapering, acuminate, 8—12 mm long, 4—8 mm broad, the petioles as long as 
or slightly longer than the blade, broadening into membranous sheath; 
cauline leaves absent; flower solitary at the summit, broadly campanulate, 

6 — 15 mm in diameter; calyx connate at bas e, the sepals lanceolate or 
elliptic, 4—7 mm long, 3—5 -nerved, as long as or longer than petals; petals 
white, with 3 light brown nerves and with numerous ferruginous dots, 
elliptic, bluntly acuminate; staminodes much shorter than petals, yellowish 
green, with 3 or 4 filiform lobes, one of which with a golden gland, the 
other shorter and eglandulose; stamens as long as petals; filaments dilated 
toward base; anthers light yellow; overy rounded -pyramidal, very light 
brown, ferruginous -puncticulate, with 4 sessile stigmas; capsule 4-lobed; 
seeds light brown with lighter border. August. 

Boggy sites in the tundra.— Arctic: Arc. Sib., Chuk. Gen. distr. : Ber., 
N. Am. Described from the shores of Lavrentiya Bay. 

3. P. bifolia Nekras. in Fl. Az. Ross., No. 11 (l 91 7) 39, Plate IV B. 
Perennial; stem glabrous, usually solitary, less often 3 or 4, stems 

10—50 cm high, faintly ribbed, slightly lustrous; leaves broadly ovate or 
oblong-ovate, entire, slightly cordate, with rounded apex or acuminate, 
1.5—5 cm long, 1—2 cm broad, paler below, 5— 7 -veined; radical leaves 4 — 12, 
with petioles longer than the blade; cauline leaves 2, at middle or on 
lower part of stem, approximate; leaves alternate, usually sessile, broadly 
lanceolate or oblong -lanceolate, amplexicaul, the lower leaf always larger 
than the upper; flower solitary, 1.5—3 cm in diameter, gaping; calyx connate 
at base, the sepals shorter than petals, 5— 11 mm long, 1— 2.5 mm broad, 
narrowly lanceolate, 3— 5 -nerved, with an apical thickening; petals obovate, 



168 



219 



9—20 mm long, 4—8 mm broad, cuneate, sometimes sinuate -margined, with 
5 — 7 very light brown nerves, ferruginous -puncticulate; staminodes shorter 
than petals, greenish or rufous, 3 -nerved, with 2 short parallel eglandulose 
lobes thickened at the tips; stamens as long as calyx, with filaments dilated 
toward base and yellowish anthers to 1 mm long; ovary brown, with 3 sessile 
stigmas, unilocular; capsule brown, 3-lobed; seeds light brown, 1 mm long. 
July — August. 

Moist meadows in the Alpine zone.— Centr. Asia: T. Sh. Gen. distr. : 
Dzu. -Kash. Described from the vicinity of Gucheng (T. Sh.). Type in 
Leningrad. 

4. P. laxmanni Pall, ex Schult., Syst. Veg. VI (1820) 696; cfr. Laxmann 
in Nov. Acta Acad. Sc. VII, 52, 241; Ldb., Fl. Ross. I, 264; Kryl., Fl. Zap. 
Sib. VI, 1438; Nekrasova, 1. c.,32.- P.turczaninowii Ldb., Fl. Ross. I 
(1842) 263.- P. ova t a Turcz., Fl. baic. -dah. I (1842-1845) 193, non Ldb. - 
Ic: Laxmann, 1. c, tab. V; Regel in Bull. Soc. Nat. Mosc. XXXIV (1861 ) 2, 
tab. VII, f. 6-9. 

Perennial, completely glabrous; rhizome short, cordlike, the roots 
slender, not dense; stem solitary, less often 2 or 3 stems, 10—20 cm 
high; leaves ovate, obtuse, with quite truncate base, rarely slightly cordate, 
cuneately tapering, entire, 0.7—3 cm long, 0.5—2 cm broad; radical leaves 
2—10, with petioles to 6 cm long; cauline leaf 1, in lower part of stem, 
sessile or short -petioled, similar in shape and size to radical leaves; 
flowers campanulate, 8—15 mm in diameter; calyx dissected to the middle 
into lanceolate, bluntly acuminate sepals, 3—5 mm long, 1—1.5 mm broad; 
petals white, 8—9 mm long, 3—4 mm broad, with 3—5 greenish nerves, obovate, 
subobtuse, tapering toward base into cuneate claw about as long as the 
blade; staminodes shorter than stamens, dissected into 3 lobes, the middle 
lobe longer than the lateral and glandiferous; filaments dilated toward base; 
anthers yellow; ovary oval -pyramidal, with 3 sessile stigmas; capsule 
light brown, trilobate, seeds light brown, to 0.5 mm long, with narrow 
membranous border. July— August. 

Boggy and moist soils on stream banks in the alpine and subalpine 
zones. — W. Siberia: Alt.; E.Siberia: Ang. -Say., Dau.; Centr. Asia: T. Sh., 
Pam. -AL, Dzu.-Tarb. Gen. distr. : Dzu. -Kash., Mong., China. Described 
from Siberia (on the way to Kamchatka). 

Note. F. subacaulis Trautv. in Bull. Soc. Nat. Mosc. XXXIII (i860) 
139.- P. sub caul is Kar. et Kir., Enum. pi. soong. 1 842, No. 140; Ldb., 
Fl. Ross. I, 773.— Plants with dense rosette of short -petioled radical leaves; 
stem small; cauline leaves very low down; flowers larger than in the 
typical form. 



Subfamily 2. HYDRANGEOIDEAE A. Br.* in Asch. Fl. Prov. Brand. I 
(1864) 61.— Woody plants with simple, mostly opposite, exstipulate leaves. 
Sepals 5, petals 5 (rarely more). Ovary half-inferior or inferior, 
3— 5-locular. Ovule with 1 integument. 



Treatment by A.I.Poyarkova. 



169 



Tribe I. PHILADELPHEAE Rchb., Consp. (1828) 43. - All flowers 
identical. Stamens usually flat. 



Genus 710. PHILADELPHUS * L. 

L. Gen. pi. (1754) 211. 

Shrubs with buds hidden under leaf traces (in our species), with opposite 
simple, dentate leaves and with large white flowers in few -flowered racemes. 
Hypanthium turbinate, adnate to ovary; sepals and petals 4; stamens 20 or 
more; ovary inferior, 3—5 -locular, with numerous pedulous ovules in each 
locule; styles 3— 5, partly connate; fruit a turbinate capsule, dehiscing 
along partitions. Seeds small, oblong, with membranous excrescence and 
fleshy endosperm. 

Economic importance. Ornamental shrubs. 

1. Flowers cream-colored, strongly aromatic; inflorescence sparse; 

young shoots bright yellow or reddish brown 

1. P. caucasicus Koehne. 

+ Flowers pure white; inflorescence dense; shoots light brown .... 2. 

2. Leaves with continuous appressed pubescence below 

*P. latifolius Schrad. 

+ Leaves quite glabrous or else the underside with beards in angles of 

veins and with solitary hairs along the veins 3. 

3. Style glabrous; leaves ovate to oval -lanceolate, sometimes very 

thin, membranous; inflorescence mostly 5 -flowered 

2. P. tenuifolius Rupr. et Maxim. 

+ Style hairy nearly to the apex; leaves usually ovate or broadly 

elliptic; inflorescence mostly 7— 9-flowered .... 3. P. schrenkii Rupr. 

1. P. caucasicus Koehne in Gartenfl. (1896) 619; Grossg., Fl. Kavk. II 
(l 930) 241 .— P. coronorius auct. fl. cauc. (non L.). 

Shrubs 1.5— 2.5m high; leaves thin, bright green above, paler below, 
oblong -elliptic, oblong -oval, or lanceolate, gradually acuminate, glabrous 
or pubescent below, with remote, inconspicuous teeth on the margin or 
entire, rarely with 8—12 developed teeth on each side; inflorescence 
„ ?1 exceeding leaves, to 14 cm long, 7—9 (ll ) -flowered, with glabrous or 

hairy awn and pedicels; flowers 2— 3.5 cm in diameter. Fl. May — June, 
fr. July— August. 

Mountain forests and forest edges, bluffs and steep slopes, to 1,800 m. 
Caucasus: Cisc. (W.), W., E., and S. Transc, Tal. (? ). Gen. distr. : 
Arm. -Kurd. (W.). Described from Abkhazia. Type in Leningrad. 

Economic importance. Nectariferous; leaves are used for the 
extraction of black dye. Wood dense; old branches used for pipe stems. 
Ornamental. 

Note. Philadelphus occurring in S. Europe — described as 
P.pallidus Hayek, ex C. K. Schn., and R. caucasicus Koehne — are 
very closely related, but should apparently be regarded as independent 
geographical races, since P.pallidus Hayek, apart from certain 
morphological differences (compact, more impoverished, 5— 7 -flowered 

* From the Greek phile in, to love and ad elphos, brother, referring to the closely approximate opposite 
shoots. 

170 



222 



inflorescence, approximate internodes and branches, darker bark, and 
firmer leaves), is further distinguished by its ecology; it is confined to shrub 
thickets on open stony slopes. The name P. coronarius L., under which 
the Caucasian and European P. pallid us are often cited, should be 
retained because Linneaus applied it to cultivated specimens of unknown 
origin belonging to different species. 

2. P. tenuifolius Rupr. et Maxim, in Bull. Acad. Sc. St. Petersb. XV 
(1857) 133.— P. coronarius Rgl. (non L. ), ibid., p. 219.— P. coronarius 
|3 tenuifolius Maxim, in Mem. Acad. Sc. Petersb., VII ser., X, 15 (1866) 
38.— P. coronarius var. satsumi Nakai, Fl. Korean. II (l 91 1 ) 485. — 

P. manchuricus Kom. in Bull. Jard. bot. Pierre le Gr. XVI (l 91 6 ) 172 
(non Nakai); Kom. and Alis., Opred. rast. Dal'nevost. kraya I (l 931 ) 610. 

Shrub 2.3 m high, with smooth or slightly hairy shoots covered by light 
brown bark, peeling in second -year shoots and replaced by gray or 
brownish-gray bark; leaves ovate to oval-lanceolate, in the forest form thin, 
chartaceous, in the form growing on open sites denser, glabrous or with 
single hairs above and with small beards in angles of veins below, with 
8—10 teeth on each side (f. d e n t at a Kom. ) or subentire (f. subinteger 
Kom.); inflorescence mostly 5 -flowered, less often 3- or 7 -flowered, 
occasionally 8— 11 -flowered (f. m ult i fl o r a Kom.); inflorescence axis 
and pedicels glabrous or hairy, flowers 2—3 cm in diameter, aromatic; 
petals oblong-oval, less often broadly oval. Fl. June, fr. August. (Plate XIII, 
Figure 1 ). 

Forests, mainly mixed and deciduous, forest edges, clearings, also in 
open sites, among stony taluses and rocks; single and in thickets. — 
Far East: Uss., Uda. Gen.distr.: Jap.-Ch. (Manch. and Korea). 
Described from the Amur. Type in Leningrad. 

Note. There are apparently 2 ecological forms : a forest form and a form 
growing on open stony slopes. The latter, apart from denser leaves, is usually 
differentiated by more compact inflorescences, with number of flowers 
often reaching 7, and by the lower growth and branching shape of the shrub. 
Morpholigically, however, these two forms cannot be separated, for 
there are imperceptible transitions between them. The open -habitat 
form has been cited for the USSR as P. manshuricus Nakai. A plant 
from the vicinity of the city of Seoul, Korea, differentiated by the dense, 
drooping pubescence of the pedicels, has been described under this name. 
Specimens similar to the latter are available from many Korean and 
Manchurian site s, but do not occur in the Soviet Union; there is therefore 
no reason to retain P. manshuricus Nakai on the list of Far Eastern 
plants. 

Economic importance. Ornamental and nectariferous plant, often 
cultivated in gardens and parks of the European part of the Soviet Union. 

3. P. schrenkii Rupr. et Maxim, in Bull. Ac. Sc. St. Pe'tersb. XV (1857) 
365; Maxim, in Mel. biol. II (1858) 542; Kom., Fl. Mansh. II (l 903) 429. - 
P. coronarius 7 satsumi Maxim, in Mem. Ac. Sc. Petersb., VII ser., 
X, 15 (l 866) 40.— P. coronarius e manshuricus Maxim., 1. c, 41 . — 
Ic: Nakai, Fl. sylv. Korean. XV (l 926) tab. XII; Kom. et Alis., Opred. rast. 
Dal'nevost. kraya I (l 931), Plate 184. 



171 



Shrub 2 3 m hig Light brown bark covering hairy shoots peeling in 
second-year shoots and replaced by brownish gray bark; leaves ovate or 
broadly elliptic, loss often oblong -elliptic, abruptly narrowed to a rather 
broad tip, with few remote minute teeth on the margin or subentire, 
loss often with well developed 10—15 teeth on each side, glabrous, or 
with scattered hairs and hairy veins below; inflorescence mostly 7-, 
loss often 5— 9 -flowered, surpassing leaves; axis and pedicels hairy, 
the pubescence sometimes present also on receptacle and sepals 
(f. c a n e s c o n s Kom.); flowers 3 — 1 cm in diameter, aromatic, with broad 
petals and hairy styles. Fl. June, fr. August. 

Forest edges and among shrub thickets, on stony slopes and among 
rook fragments: growing singly, rarely in thickets.— Far East: Uss., 
Uda. Gen. distr. : Jap.-Ch, (Manchuria, Korea). Described from the 
village of Pakhale on the Amur. Type in Leningrad. 

Economic importance. Ornamental and nectariferous plants. Cultivated 
in gardens and parks of the European part of the Soviet Union. 

Note. The specific independence of P. s c h r en ki i is very doubtful: 
it is very closely related to P. t e nui f oli u s and, in practice, cannot 
always be differentiated from specimens of the latter growing on open 
slopes; the only distinctive character of P. schrenkii — pubescence of 
the style — varies greatly, and some specimens have very few hairs 
at base of style; this may sometimes be observed also in the forest form 
of P. tenuifolius. 

*P. latifolius Sehrad. ex DC. Prodr. Ill (1828) 206. 

Shrub, growing profusely, with light yellow shoots; leaves densely 
gray -hairy below, on fruiting shoots oval or oblong, entire or minutely 
denticulate, on sterile shoots rounded -ovate, with large teeth; flowers 
large, white, aromatic, in 5— 7 -flowered racemes. Fl. June — July, 
fr. September. 

N. Am. (Tennessee) — rocky riverbanks. Often cultivated in the European 
part of the Soviet Union, to Leningrad inclusive. 

Note. Apart from this species, the following foreign species are most 
often cultivated : the European P. pallid us Hayek and the N. American — 
P.pubescens Loiss, a species closely related to P. la t i f o 1 i us, but 
distinguished by its smaller leaves, lower growth, and more nearly closed 
flowers; P.inodorus L., with smooth light brown shoots, broad, 
subglabrous, entire or slightly dentate leaves, and large white, almost 
odorless flowers with stigmas longer and broader than anthers. 



Genus 711. DEUTZIA THBG. 
rhbg.,Nov.gen.pl.I (1781) 19. 

Shrubs with opposite, simple, dentate leaves and a pubescence of 
unicellular stellate hairs. Flowers white, rather large, in compound 
corymbiform or oblong racemes; hypanthium campanulate, adnate to ovary. 



. after the Dutchman Van Deutz, a companion of Thunberg. 



5773 172 



Lignifying in fruit; sepals and petals 5; stamens 10, rarely 12— 15, with 
flat, often apically tripartite filaments; ovary inferior, 3- or 4-locular, 
with many ovules in each locule; styles 3 or 4; fruit a subglobose 
capsule, dehiscing below along partitions into separate carpels, the 
latter dehiscing at the apex and remaining united dorsally. Seeds small, 
with a seed -coat, tubularly broadening below, with a small wing at the apex. 

224 1. Flowers ca. 1 cm indiameter,the hypanthium gray with uninterrupted 

stellate pubescence. Capsule 2 mm long, 2.5 mm broad, retaining dense 

pubescence 1. D. amurensis (Rgl.) Airy -Show. 

+ Flowers 1.5 cm in diameter, with glabrous hypanthium; capsule 3 mm 

Long, 3.5—4 mm broad, glabrous 2. D. glabrata Kom. 

1. D. amurensis (Rgl.) Airy-Show in Bull. Misc. Inform. (1934) 179. — 
\). p a r v i f 1 o r a Maxim., Prim. fl. amur. (l 859) 110, non Bge. ex parte; 
Kom., Fl. Mansh. II (1903) 431.- D. parviflora var. amurensis Rgl., 
Tent. Fl. ussur. (1861) 62. — ? D. c o ry mbo s a var. parviflora C.K. Schn., 
Illustr. Handb. Laubh., I (1906) 382.- Ic: Rgl., I.e., tab. 5, f. 14; Nakai, Fl. 
sylv. Koreana XV (1926) tab. XX.- Exs.: HFR, No. 1928. 

Small spreading shrub; curved young shoots covered with light 
brown bark later turning gray; leaves oval-elliptic or oblong-elliptic, 
finely serrulate, glabrous below, with remote stellate hairs above; 
inflorescence corymbiform, the axes and pedicels more or less densely 
covered with only stellate hairs; filaments subulate or obscurely dentate 
at the apex. Fl. June, fr. August. (Plate XIII, Figure 2). 

Mixed forests and shrub thickets.— Far East.— Uss. Gen. distr.: 
Jap. -Ch. (Manchuria, N. Korea). Described from the Amur. Type in 
Leningrad. 

Economic importance. Ornamental plants. 

2. D. glabrata Kom., Fl. Mansh. II (1903) 433.- D.glaberrima Koehne 
in Engl. Bot.Jahrb.XXXIV, Beibl. LXXV (1904) 308.- D. fa uriei Lev. in 
Fedde Repert. VIII (1910) 283.- Ic: Nakai, Fl. sylv. Koreana XV (1926) 
tab. XXI. 

Spreading shrub to 2 m high, with angular or curved branches; reddish 
brown bark covering young glabrous shoots, peeling later and replaced 
by S ra y or brownish gray bark; leaves oblong-elliptic or lanceolate, with 
cuneate base, tapering to a more or less short mucro, regularly serrulate, 
glabrous below, with scattered stellate hairs above; flowers a loose 
corymbiform panicle with glabrous axes and pedicels; filaments tapering 
upward, with no excrescences or teeth. Fl. June, fr. August. 

Stony forest edges, near and on rocks, always in the shade.— Far East: 
Uss. — only one habitat near the village of Bidzhanskoe on the upper 
Lugovaya River. Gen. distr.: Jap. -Ch. (Manchuria and Korea). Described 
?2 from the Chari-Kori Valley in N.Korea. Type in Leningrad. 

Note. The Japanese D. s c ab r a Thunb. (D. c r e n at a S. et Z.) is 
often cultivated in southern districts of the Soviet Union; all its parts — 
including both sides of leaves — densely covered with stellate hairs; 
flowers in oblong, erect racemes, the filaments tripartite at the apex. 



173 



Tribe 2. HYDRANGEAE DC.Prodr. IV (1830) 13, p. p.- Outer flowers 
often Sterile, with large calyx; stamens filiform or subulate. Fruit a 
capsule or berry. 



Genus 712, hydrangea L. 

I,. Sp.pl. i I 753) 397. 

Shrubs or trees with opposite, simple, dentate leaves; inflorescence 
a large, corymbiform panicle, the outer flowers sterile, composed of 
3—5 large, pelallike sepals, the other flowers small, bisexual. Sepals 
and petals 4 or 5. Petals sometimes connate at the apex. Stamens 8—10 
or 15—20. Ovary inferior or half-inferior, with numerous ovules, 
2— 5-locular; styles 2—5, short, free or connate at the base. Fruit a 
I - or 5-locular capsule dehiscing at the apex. Seeds small, often winged. 

1. Leaf blade more or less equaling petiole, 4— 10 cm long; petals 
connate at the apex, falling at opening of flower; ovary inferior; 
seeds not winged 1. H. petiolaris S. et Z. 

+ Leaf blade several times longer than petiole, the latter 1—3 cm long; 
petals free, persistent until pollination; ovary half-inferior; seeds 
winged 2. S. paniculata Sieb. 

1. H. petiolaris S. et Z., Fl. jap. I (1835) 106. - H. c or dif olia S. et Z., 
1. c, 113. — H. s c a nd en s Maxim, (non DC.) in Mem. Acad. Petersb., ser. 
VII, XII (1868) 130.- Ic: S. et Z., 1. c, tab. 54, 59. 

Climbing shrub, ascending to 3—6 m, with glabrous, reddish brown, 
rooting shoots; leaves thin, glabrous above, with beards confined to angles 
of main veins below, broadly oval or broadly elliptic, with rounded, broadly 
cuneate or shallowly-cordate base, the margin regularly and acutely 
denticulate; inflorescence glabrous, corymbiform, to 18 cm in diameter; 
stamens 15; sepals of outer flowers entire or more or less dentate. 
Fl. July— August, fr. September. 

Deciduous and coniferous forests and subalpine mountain zone. — 
Far East: Sakh., known in southern part of island; occurrence in northern 
part doubtful. Gen. distr.: Japan (Hokkaido, Honshu, Kyushu). Described 
P ?r from Japan. 

Economic importance. Ornamental plants. 

2. H. paniculata Sieb. in Nova Acta Acad. Leop. XIV, 2 (182 9) 691; 

Fr. Schmidt, Fl. sachal. in Mem. Acad. Petersb., ser. VII, XII (1868) 130.- 
Ic: S. et Z., Fl. jap. I (1835) tab. 61; Miyabe et Kudo, Ic. forest, trees 
Hokkaido, tab. 46. 

Erect, branching shrub to 6 m high, with glabrous and slightly hairy 
reddish -brown shoots; leaves firm, broadly oval, less often oblong -oval 
or elliptic, dark green, scattered -hairy above, light below, with dense, 
appressed hairs along the veins and scattered hairs throughout the surface, 



From the Greek hydro, water and angcion, vessel - the fruit is shaped like a bowl. 



174 



with broadly cuneate base; inflorescence pyramidal, densely hairy; 

stamens 10; sepals of outer flowers entire. Fl. July — August, fr. September. 

Riparian and mountain forests; tolerating boggy soils.— Far East: Sakh., 
until now known in southern part of the island as far as the Poronai River. 
Gen. distr. : Japan. Described from Japan. 

Economic importance. The bark is used for the manufacture of Japanese 
paper; the wood, which is very hard and white, is used for various small 
articles. Ornamental plant, often cultivated in the European part of the 
Soviet Union. 

Note. Among the species of this genus represented in the USSR, the 
one most cultivated isH. opuloides C. Koch. (Hortensia opuloides 
Lam., Hy d r a ng e a hortensis Sieb.) — originally from Japan, with 
glabrous shoots, elliptic mucronate leaves cuneately tapering toward the 
base, and large inflorescences; the flowers of the garden forms are all 
sterile (white, pink, or sky blue). Cultivated in northern regions in flower pots, 
in southern regions in the ground. Escaped on the Caucasian coast and 
sometimes forming thickets (near Batumi). 



Subfamily 3. RIBESIOIDEAE Engl.- in Engl, et Pr. Nat. Pflzf. Ill, 2a 
(1890) 88.— Woody plants with simple alternate exstipulate leaves. Flowers in 
racemes. Petals 5. Ovary unilocular, with 2 parietal placentas. Fruit a berry. 



Genus 713. RIBES *# L. 

L. Gen.pl. (1737) 68. 

Shrubs with smooth or spiny shoots, with alternate, palmatilobate, 
dentate leaves and with flowers in simple racemes; pedicels developed; 
flowers bisexual or dioecious, 5-merous; hypanthium from flat to tubular, 
adnate to ovary below, passing directly into sepals at the apex; petals much 
smaller than sepals; stamens 5, opposite to sepals, attached to margin 
of hypanthium or slightly lower; styles 2, mostly connate to above the 
middle; ovary bilocular, smooth or glandular, inferior, very rarely half- 
inferior, short -stalked. Fruit a succulent berry with dried perianth at 
the apex, at maturity disarticulating from pedicel; seeds with interior 
hard endopleura and gelatinous testa. 

1. Shoots densely covered with acicular spines and, in addition, with 
larger spines at the nodes, 7—15 in each verticillaster; leaves 
rigid-hispid; flowers flat; berries black, glandular-hispid. Subgenus 
Grossularioides Jancz.) 15. R. horridum Rupr. 

+ Shrubs unarmed or with short paired spines at the nodes, sometimes 

also with spiny internodes; leaves not hispid 2. 

2. Leaves with yellow punctate aromatic glands below. (Subgenus 
Eucoreosma Jancz.) 3. 

+ Punctate aromatic glands absent 13. 

3. Flowers campanulate, tall, erect shrubs. (Series Nigra e) 4. 

+ Flowers flat, cuplike. Mostly low shrubs 10. 

" Treatment by A.I. Poyarkova. 

* From the Arabic Ribas, as rhubarb (Rheu m ribes L.) was called by the Arabs in their own country. 

After their penetration into Spain, they transferred this name to the acid fruit of the currant growing 

in that country. 

175 



228 



4. Hypanthium short, 1.5—2 times broader than high 5. 

+ Hypanthium higher, as high as or higher than broad 7. 

5. Flowers pale, usually yellowish, rarely flesh-colored, with spreading 

or slightly recurved sepals 6. 

+ Flowers lilac or pinkish gray, the sepals recurved and often appressed 
to receptacle 21. R. nigrum L. 

6. Sepals spreading; hypanthium slightly pentagonal; ovary high-conical; 
leaf lobes acute; pedicels slender, nodding . . . 16. R. ussuriense Jancz. 

+ Sepals suberect or divaricate; hypanthium not angled, smooth; leaf 

lobes broad, mostly subobtuse; pedicels thick 

17. R. pauciflorum Turcz. 

7. Flowers large, 10 — 12 mm long, flesh-colored, broadly campanulate; 
leaves large, to 15 cm in diameter, glabrous, lustrous, deeply and 
acutely lobate 20. R. janczewskii A. Pojark. 

+ Flowers 8—9 (lO)mm long, turbinate or cyathiform 8. 

8. Hypanthium constricted above, turbinate, pentagonal; leaves firm, 
lustrous, acutely dentate 19. R. turbinatum A. Pojark. 

+ Hypanthium cyathiform, its height 1.5—2 times as great as or equal 

to width; leaves dull 9. 

9. Hypanthium 1.5—2 times as high as broad; styles split to the middle 

or deeper; leaves mostly with simple large teeth . 

18. R. kolymense Kom. 

+ Hypanthium as high as or 1.5 times higher than broad; style entire 

or split at the apex; leaves duplicato-dentate 

21. R. nigrum var. sibiricum E. Wolf. 

10. Tall shrub with large, cordate, deeply and acutely lobate leaves; 

berry blue 22. R. dikuscha Fisch. 

+ Low shrubs with small (2—5 cm in diameter), reniform or rounded - 

reniform leaves with slightly developed lobes; berries brown .... 11. 

11. Leaves tomentose -pubescent below 25. R. graveolens Bge. 

+ Leaves with only punctate glands below 12. 

12. Leaves densely glandular below, very aromatic; flowers white 

24. R. fragrans Pall. 

+ Leaves with sparsely scattered glands below; flowers purple 

2 3. R. procumbens Pall. 

13. Flowers bisexual 14. 

+ Flowers dioecious. (Subgenus Berisia Spach) 31. 

14. Flowers large, 10—15 (20)mm long, with high campanulate -tubular or 
cylindrical hypanthium, deep pink or yellow 30. 

+ Flowers small, to 6 mm long, with low patelliform or short -campanulate 
hypanthium, usually greenish, often more or less dingy purple, rarely 
white or pinkish 15. 

15. Prostrate, low shrubs with white or pinkish flowers and glandular - 
hispid ovaries and berries (Subgenus Heritiera Jancz.) 16. 

+ Erect, unarmed, mostly tall shrubs with greenish or more or less 
dingy purple flowers and smooth ovaries and berries. (Subgenus 
Ribesia Jancz.) 17. 

1 6 . Leaves to 9.5 cm broad, deeply 5 - or 7 -lobed 

13. R. sachalinense Nakai. 

+ Leaves to 3.5 (4) cm broad, with slightly developed lobes 

14.. R. malvifolium A. Pojark. 



176 



229 



230 



17. Hypanthium cuplike or campanulate, with smooth base 19. 

+ Hypanthium quite flat, patelliform, with fleshy perigynous ring at 

the base. (Series Vulgares) 18. 

18. Flowers dingy purple or, at least, with colored hypanthium and petals; 
stamen connectives much narrower than anthers; bark dark brown, 
separating in large lamellate strips .....' 1 . R. triste Pall. 

+ Flowers greenish; stamen connectives broad, as long as anthers; 

bark gray, peeling *R.vulgare Lam. 

19. Hypanthium shallow, cuplike; sepals glabrous on the margin or 
sparsely ciliate 20. 

+ Hypanthium campanulate or turbinate; sepals with dense border of 

cilia 25. 

20. Sepals reflexed nearly from base, hence stamens strongly excerted; 
leaves with acute lobes, the middle lobe usually much longer than the 
lateral (Series Mul t i f 1 o r a e) .... 2. R. manschuricum (Maxim.) Kom. 

+ Sepals with spreading limb; only anthers protruding; upper leaf lobes 
usually more or less equally developed. (Series Rubrae) 21. 

21. Annual shoots light brown; leaves tetragonal in outline with truncate 
base, usually hairy only along the veins below; racemes short, erect, 
dense, with densely hairy axis and pedicels; flowers pale, small . . 
3. R. palczewskii A. Pojark 

+ Shoots pale yellow; raceme loose, glabrous or with sparse 

pubescence 22. 

22. Racemes long, to 9 (ll) cm long, with up to 22 flowers, horizontal or 
nodding; sepals usually with brown-red spots and veins, sparsely 
ciliate-margined; leaves pubescent below 6. R. pubescens Hedl. 

+ Racemes shorter, erect at anthesis; sepals greenish, sometimes with 
purple spots or brownish, glabrous on the margin 23. 

23. Racemes short, 2 — 5 cm long, 3 — 8 (10 )-flowered, loose; flowers to 

6 mm in diameter; leaves mostly glabrous, lustrous, with acute lobes, 

coarsely dentate 4. R. rubrum L. 

+ Racemes 6 — 12 -flowered; flowers smaller, 3 — 4 (5) mm in diameter 

24. 

24. Flowers pale, small, usually not more than 3 — 3.5 mm in diameter; 
inflorescence axis, pedicels, young shoots, and petioles usually more 
or less glandular-hispid, with an additional simple pubescence; leaves 

mostly pubescent below, broad, with short obtuse lobes 

5. R. hispidulum A. Pojark. 

+ Flowers brownish, larger; leaves with acute, mostly well developed 

lobes *R- scandicum Hedl. 

25. Flowers turbinate, with erect sepals; styles cylindrical; berry black, 
with purple-lilac sap. (Series M e y e r i an a e) .... 7. R. meyeri Maxim. 

+ Flowers campanulate; sepals with recurved or spreading limb; style 
conical; berry red or purple-black. (Series Petraeae) 26. 

26. Bark on shoots light brown, separating in large lamellate strips; 

flowers yellowish brown; berry black with purple sap 

10. R. altissimum Turcz. 

+ Bark light gray, peeling; flowers purple 27. 

27. Hypanthium with fleshy excrescences below the petals; flowers purple; 
berry dark red or cherry-red 12. R. biebersteinii Berl. 

+ Hypanthium without excrescences; berry red 2 8. 

28. Flowers pale, yellowish or pinkish, with narrow, conical styles; berry 
light red 9. T. pallid if lorum A. Pojark. 

177 



+ Flowers dark purple; berry vivid red 29. 

29. Ovary apex in the shape of a low cone; flowers large, 6 — 7 mm long; 
leaves to 15 cm broad 8. R. latifolium Janez. 

+ Ovary apex high-conical; flowers 4 — 5 mm long; leaves usually not 

more than 10 cm in diameter 1 1 . R. atropurpureum C A. M. 

30. Flowers dark pink; ovary glandular-hairy; berry glandular; leaves 
whitish-tomentose below *R. sanguineum Pursh. 

+ Flowers bright yellow; ovary, berry and leaves glabrous 

*R. aureum Pursh. 

31. Pubescence of appressed or crisp hairs, with all glands stalked; 
flowers and berries glabrous 35. 

+ Pubescence of spreading, straight white hairs; in addition to the 

stalked glands, there are often also sessile, viscous glands; flowers, 
and often also berries, hairy. (Series Orientales) 32. 

32. Flowers pale greenish; leaves with well developed lobes, dark green, 
lustrous, usually with bristly glandular hairs above; berries red . . 
26. R. orientale Desf. 

+ Flowers purple; leaves without glandular bristles above 33. 

noi 33. Leaves 1—2 cm broad, thin, cordate, with well developed lobes; shoots 

slender, pale yellowish gray; viscous glands absent 

27. R. melananthum Boiss. et Hoh. 

+ Leaves to 3 — 3.5 cm broad, firm, with slightly developed subobtuse 

lobes; shoots light brown; viscous glands usually present 34. 

34. Leaves and shoots densely hairy, usually with few viscous glands; 
shoots thick 2 9. R. villosum Wall. 

+ Leaves glabrous or slightly hairy, with slight addition of viscous 

glands; shoots slender 28. R. heterotrichum C. A. M. 

35. Leaves trilobate or obscurely 5-lobed, with well developed upper 

lobes 37. 

+ Leaves incised in upper part into 3 slightly developed, obtuse lobes; 
shoots light brown, usually with a pair of spines at base of leaves, the 
sterile shoots sometimes with spiny internodes. (Series Diacanthae) 
36. 

36. Leaves rounded-cuneate, dull, usually puberulent below; inflorescence 

axis and pedicels pubescent; berries dark cherry-red 

35. R. saxatile Pall. 

+ Leaves oblong-cuneate, quite glabrous, lustrous; inflorescence 

glabrous; berries red 34. R. diacantha Pall. 

37. Shoots light brown, lustrous, usually with a pair of spines at base of 
leaves; leaves small, 0.3 — 3.2 cm long. (Series P u 1 c h e 1 1 a e) . . . 
36. R. pulchellum Turcz. 

+ Shoots dull; gray or pale brown, unarmed; leaves to 4.5 — 5 cm 

broad 38. 

38. Fruiting racemes short, 3 -flowered; leaves dull, usually covered with 
glandular bristles above, mostly with cordate base. (Series A 1 p i n a e) 
39. 

+ Fruiting racemes (4) 5 — 1 1 -flowered; leaves lustrous, mostly glabrous 
above, with cuneate or truncate base. (Series Lucidae) 40. 

39. Berry oblong, oval or obovate; leaf lobes acuminate, the upper lobe 
protruding, rhomboid 30. R. maximowiczianum Kom. 

+ Berry globose; leaf lobes more or less equally developed, mostly 

broadly triangular, acute 31 . R. alpinum L. 



178 



40. Leaves orbicular, with slightly developed, obtuse lobes; fruiting 

racemes 6 — 11 -flowered 32. R. komarovii A. Pojark. 

+ Leaves oblong-cuneate, with strongly protruding middle lobe; 

fruiting racemes (4) 5 — 8-flowered 33. R. lucidum Kit. 

Subgenus 1 ." RIBESIA (Beri.) Jancz. in Bull. Acad. Cracov (1903) 
236; Berl. in Mem. Soc. phys. Gen. Ill, 2 (1826) 43, pro sect. — Flowers in 
racemes, bisexual, greenish or purple, with flat, campanulate or cylindrical 
hypanthium and smooth ovaries. Glands stalked, crystalline; bud scales 
coriaceous light brown. 

Economic importance. The species of this subgenus are the ancestors 
of cultivated red currants, the separate varieties either originating directly 
from or being hybrids of wild species. The berries of all wild species 
are edible and differ from cultivated currants mainly in their sour taste, 
some also in their coarse, thick skin. 



Series 1. Vulgares A. Pojark.— Hypanthium flat, patelliform, with broad 
base, with fleshy pentagonal, perigynous ring; sepals reflexed, glabrous. 
Stamens and styles short; berry red. 

1. R. triste Pall, in Nova Acta Acad. Petrop. X (1797) 378 (excl. pi. 
fruct.); Ldb., Fl. Ross. II, 1, 198; sensu stricto: Maxim, in Mel. biol. IX 
(1873)235; Kom., Vved. v izuch. rast. Yakutii.I (1926) 142; Fl. Kamch. II 
(1929) 225; Kom. and Alis., Opred. rast. Dal'nevost. kraya I (1931) 621.- 
R. vulgaris fr. rubro Krascheninnik., Beschr. Kamtsch. (1766) 101 . — 
R. inerme floribus planiusculis Gmelin, Fl. Sib. Ill (1747) 173. — 
R. m elanc hoi i c um Sievers ex Pall., I.e., — R. p r o p i nquum Turcz. in 
Bull. Soc. Nat. Mosc.XHI (1840) 70; Ldb., Fl. Ross. II, 199.— R. rub rum j3 
propinquum Trautv. et Mey., Fl. ochot. (1856) 40.— R. rubrum var. 
rubellum Rgl. et Til., Fl. ajan. (l 859) 92. — R. r u b r um y subglandu- 
losum Maxim, in Mel. biol. IX (1873) 234.— R. rubrum var. glabella 
Trautv. et Mey. in Syll. pi. sibir. (1888) 32; Trautv. et Mey., Fl. ochot. 
(1856) 40.— Ic: Jancz. in Monograph, d. Groseill. (l 907) f. 24; Poyarkova 
in Tr. Bot. Inst. Akad. Nauk SSSR, ser. 1, No. 2 (1936) Figure 1 (p. 163). 

Shrub with mostly decumbent branches and numerous, scarcely 
branching shoots, ascending to 20 — 40 cm, less often erect, to 75 cm high, 
the light brown bark separating in large strips; leaves broad, 3 — 5-lobed 
acutely and coarsely dentate, usually glabrous, less often pubescent below, 
to 6.5 cm long and 9 cm broad; raceme short, loose; pedicels 2.5 — 5 mm 
long, slender, pubescent; petals mostly broad, flabellate, rarely narrower, 
cuneate, U~ lz as long as sepals; style thick, split nearly to base; berry 
bright red, 6 — 10 mm in diameter, very succulent. FL June, fr. August. 
(Plate XIII, Figure 5). 

Banks of rivers and streams, riverain deciduous forest strips, also 
slopes, mainly rocky slopes, shrub thickets and forests, rarely bogs; 
solitary or in groups.— Arctic: Arc. Sib., An.; E. Siberia: Lena-KoL, 
Dau.; Far East: Kamch., Okh., Ze.-Bu., Uda, Uss. Gen. distr.: Jap. -Ch. 
(N. Korea), N. Am. (Canada and Northern states). Described from Yablonovyi 
Range. Type in London. 



179 



Economic importance. The berries are quite edible. The plant may be 
used for selection and hybridization with cultivated varieties of red 
currants for the breeding of cold -resistant varieties for northern regions. 

Note. Japanese and American plants have been described as separate 
varieties (R. r u b r u m var. bracteosum Maxim. —Japanese, R.triste 
var. a lb in e r v ium (Mchx.) Fernald. — R. a lb in e r v ium Mchx. and 
R.triste var. a la s ka num Berger.— American). Attempted clarification 
of the taxonomic significance of these forms has been unsuccessful owing 
to lack of material. 

*R. vulgare Lam., Encycl. met. bot. Ill (1789) 47. — R.sativum Syme 
in Engl., Bot. Ill, ed. IV (1865) 42. — R. domesticum Jancz. in Compt. 
rend. XXVI (1900) 588.- R. s ilvestre Hedl., Bot. notis. (1901) 82.- 
R.hortense Hedl., 1. c. — R. r u br u m (ex parte) Shmal'g., Fl. I, 358 
(non L.). — R.rubrum auct. plur. (non L.). — Ic. : Jancz., Monogr. d. gros. 
(1907) f. 18-21. 

Erect shrub with light grayish shoots; leaves 3 -or 5-lobed, with short, 
acute lobes, coarsely dentate, glabrous on both sides, less often hairy 
below, shallowly cordate, racemes long, to 8 cm, loose; axis and pedicels 
glabrous, 3—5 mm long; flowers greenish; style slender, split to the 
middle or less. Fl. May —June, fr. July— August. (Plate XIII, Figure 3). 

Cultivated for its berries, but apparently not escaping. Grows wild 
in the mountains of France, in southern Belgium, N. Italy, and the 
Pyrenees. Described from France. Type in Paris. 

Note. 1. R. vulgare Lam. is the ancestor of most garden varieties 
of red currants, which originated from it either directly or by way of 
crossing with other species of the subgenus R i b e s i a, apparently with 
R.petraeum Wulf and R. p ub e s c e ns Hedl. Var. ma c r o c a r p um 
Jancz. is distinguished from the typical form by its larger flowers and 
fruits, and by its peculiar shape, resulting from the fact that only the buds 
arranged at the base of shoots reach a complete development, while the 
higher buds die off together with the shoots bearing them. 

Note 2. In the Soviet literature, R. v u 1 ga r e Lam. is usually cited 
under the incorrect name of R.rubrum L. and is confused with 
R. pubescens Hedl. (e. g., in the floras of Syreishchikov and Maevskii), 
which grows wild in the European part of the Soviet Union. 

Economic importance. The berries are used both raw and for the 
„„ . making of jam, syrup, jelly, wine, vinegar, and refreshing fruit drinks. 

According to Hegi, the berries contain the following percentages: 83—85.8 
water, 4— 9 invert sugars, 1.84—2.59 free acids (especially citric acid), 
1.47 pectins, pectose, 0.41 pentosans, 0.35— 0.70 albumins, ca. 4 -cellulose, 
0.72 -ash, red pigment (cyanidin-glucoside); the seeds contain 16—18% fatty 
oils. The red pigment of the berries colors linen yellow. 



Series 2. Multiflorae A. Pojark. — Hypanthium cuplike, with fleshy 
excrescences below petals; sepals recurved nearly from base, glabrous 
on the margin or sparsely ciliate; style narrowly conical; ovary inferior; 
berry red. 

Apart from our species, R.multiflorum Kit., distributed in the 
mountains of southern Europe, also belongs to this series. 



180 



235) 




PLATE XIII. 1-PhilaHpl h 

A^-show, a) fruit; 3_ R ,r; u :2;:r;:i e r m - ; — utziaa amurensis(Rg] 

-- Pan., ) flower from above> ^fliSSi.T? ° f f ^" ^ ^^escens utl 
11 Max ^-.fiower, a) flower section R-vul gare Lam., flower; 7- 



A 

5 - R.t 
R.meye 



181 



2. R. manshuricum (Maxim.) Kom., Fl. Mansh. II (1903) 4 37; Kom. and 
Alis., Opred. rast. Dal'nevost. kraya I (1931 ) 621.— R.multiflorum 
var. manshuricum Maxim, in Mel. biol. IX (1873) 228.— R.petraeum 
var. mongolicum Franch. in Nouv. Arch. Mus. Paris. 2 ser., VI (1883 -1 884) 
7. - Ic. : Kom. and Alis., 1. c, Plate 185. 

Shrub 1—2 m high; shoots light, the bark separating in small, thin lamellae; 
leaves deeply cordate, mostly trilobate, with long-acuminate lobes, the 
middle lobe usually much larger than the lateral, glabrous above, with sparse 
hairs along the veins below (f. s ub gl ab r u m Kom.), or scattered -hairy 
above and pubescent below (f. villosum Kom. ); racemes 2 . 5—9 cm long, 
rarely to 16 cm long, dense, spiciform, with up to 40 flowers; pedicels 
1—2 mm long; inflorescence axis thick, pubescent; flowers greenish, with 
much exserted stamens; berry 7 — 9 mm in diameter, very acid, with thick 
skin. Fl. June, fr. August. (Plate XIV, Figure 1 ). 

Forest edges, clearings in mixed forests; solitary. In rock gorges 
in the shade of rocks, forming small thickets.— Far East: Ze.-Bu., Uda, 
Uss. Gen. distr.: Jap.-Ch. (Manchuria, Chihli Province, N. Korea). 
Described from the Chaorie Pass in N. Korea. Type in Leningrad. 



Series 3. Rubrae A. Pojark. — Hypanthium flat, cuplike, smooth inside; 
sepals with spreading limb, smooth on the margin or sparsely ciliate; 
9 on style cylindrical; ovary inferior, with slightly conical apex; berries red. 
Apart from our species, R.spicatum Robs. — from the British Isles 
including Ireland — also belongs to this series. 

Economic importance. The berries of all species of this series may 
be used like those of the cultivated red currant R.vulgare Lam., from 
which they are distinguished by their more acid taste and coarser skin. 

3. R. palczewskii A. Pojark. in Bull, of Appl. Bot. Genet, and Plant Breed 
XXII, 3 (1929) 341; Acta Inst. Bot. Ac. Sc. URSS, Ser. 1,2 (1936) 168; 
Kom. and Alis., Opred. rast. Dal'nevost. kraya I (l 931 ) 622.— R.rubrum 
a g lab e Hum Trautv. et Mey., Fl. ochot. (1856)40.- R.rubrumy 
acuminatum Rgl. in Mem. Acad. St. Petersb. VII ser., IV, 4 (1861 ) 67.- 
R. rub rum (3 silvestre Maxim, in Mel. biol. LX (1873) 233 (ex parte). - 
R. rub rum var. sativum Korsh. in A. H. P. XII (1892) 340.- 
R.pubescens (non Hedl.) Kom., Fl. Mansh. II (1903) 441.- R. rub rum 
var. palczewckii Jancz. in Monogr. d. grosseill. (1907) 290. — 
R.warscewiczii (non Jancz.) Kom. and Alis., Mai. opred. rast. D. Vost. 
(l 925) 251 . - Ic. : Kom. and Alis., Opred. rast. Dal'nevost. kraya I (l 931 ), 
Plate 187; Poyarkova in Tr. Bot. Inst. Akad. Nauk SSSR. ser. 1, No. 2 (1936), 
Fig. 3 (p. 167). 

Shrub ca.l— 1 .5 m high; shoots glabrous or slightly glandular, with smooth, 
light brown bark; leaves broad, glabrous above, pubescent along the veins below, 
rarely glandular -hairy along the veins or uninterruptedly pubescent, 3 - or 
5-lobed, with short, broadly triangular lobes, shallowly cordate or truncate 
at base; racemes erect, 2— 5 cm long, 5— 15 -flowered, dense; axis and 
short (l— 2.5 mm) pedicels more or less densely lanate-pubescent; flowers 
small, 3—3.5 mm in diameter, yellowish; berry to 8 mm long, mostly oblong. 
FL end of May, June, fr. July. 



182 



Forests and riverside shrub thickets.— E.Siberia: Lena-Kol. (S. part — 
slopes of the Yablonovyi Range), Dau. (E. part); Far East: Ze.-Bu., Uda, 
Uss. (N. part — lower reaches of the Amur and Ussuri rivers). Gen.distr.: 
unknown outside the regions mentioned; probably grows also on right 
(Chinese) bank of the Amur. Described from the village of Voskresenskoe 
on the Amur River. Type in Leningrad. 

Economic importance. Deserves attention as a baccate plant. 

4. R. rubrum L., Sp.pl. (1753) 200; Turcz., Fl. baic. -dah. I, 443; Ldb., 
Fl. Ross. II, 1, 199 (ex parte). — R. i ne r me floribus planiusculis 
Gmel., Fl. sib. Ill (1747) 173. - R. ac idum Turcz. in Ldb., Fl. Ross. II, 1, 
199 (pro synon. R. r ubr i); Poyarkova in Tr. Prikl. Bot., Gen. i Sel. XXII, 3 
(192 9) 341.— R.baicalense Turcz., 1. c. — R. rubrum var. silvestris 
Trautv., PL Czekanovsk. et Mill. (1877) 57.- R. glabellum Hedl. in Bot. 
Notis. (1901) 98; Komarov, Vved. v isuch. rast. Yakutii I (1926) 143.- 

R. rubrum var. g 1 ab e 1 lum Jancz., Monogr. d. groseill. (l 907) 289. — 
R. rubrum var. s c a n d i c u m (non Jancz.), Fedch. i Fler., Fl. Evr. Ross. 
(1910) 504.- Ic: Poyarkova in Tr. Bot. Inst. Akad. Nauk SSSR, ser. I, No. 2 
(1936), Fig. 5 (p. 171). 

Shrub to 1 m high; shoots glabrous or more or less beset with glandular 
hairs, covered with smooth pale yellow bark; leaves usually deeply 
cordate, 3- or 5-lobed, with acute, coarsely dentate lobes, mostly glabrous 
on both sides, lustrous, less often pubescent below or glandular -hispid 
along the veins; petioles glabrous or glandular -hispid, often reddish; 
racemes erect, 2—5 cm long, loose, 3—8 (lO)-flowered, nodding in fruit; 
axis and long (5—12 mm) pedicels glabrous or glandular; flowers rather 
large, greenish, often with purple petals and macular sepals; berry 8—11 mm 
in diameter, sometimes oblong. FL May — June, fr. July —August. (Plate XIII, 
Figure 3). 

Forests and forest edges, riverain deciduous forest strips, river slopes, 
shrub thickets. — Arctic : Arc. Eur., Arc. Sib. (W. part); European part : 
Kar. -Lap. (N.), Dv.-Pech. (N.), Urals (as far South as 60°lat.); W.Siberia: 
Ob, Alt. (N. in subalpine zone); E.Siberia: Yenis., Lena-Kol. , Ang. -Say., 
Dau. (W. part). Gen.distr.: Mong. (mountains of N. and Centr. Mong.). 
Described from northern Sweden. Type in London. 

Note. The distribution area of R. r ubr um covers the northern part 
of Scandinavia, the northern part of the European territory of the Soviet 
Union, and the Arctic part of Siberia (as far east as the Kotuiand Mon'ero 
rivers), the southern part of Yakutia (to the Vilyui River), and E. Siberia 
from the Yenisei to W. Transbaikalia; replaced in W. Siberia by 
R.hispidulum. Hybrids of R. rubrumX hispidulum occur very 
frequently in the Yenisei area. 

Economic importance. Combines the high quality of its fruit with good 
winter hardiness; thus it deserves the attention of fruit-growers for 
experimental breeding of red currant varieties suited to northern regions 
of the Soviet Union. 

5. R. hispidulum A. Pojark. in Bull, of Appl. Bot. Genet, and Plant. Breed. 
XXII (1929) 339; Kryl., FL Zap. Sib. VI (1931) 1440.- R. rubrum a et |3 

C. A. M. in Ldb., Fl. Alt. I (1829) 267. - R. r ub r u m (ex parte) Ldb., Fl. Ross. 
11,1,199.— R. rubrum var. glabella Trautv. in Bull. So c. Nat. Mo sc. 



183 



239 



240 



XXX K, 1 (1866)315 (non Trautv. et Mey.). - R. pube s c e ns Kryl., Fl. 
Alt. I (l 903) 4 64 (non Hedl.). — R.rubrum ssp.asiaticum Jancz. ex 
C.I. Schn., Illustr.Handb.d. Laubh. I (1906) 430 (ex parte?).- R. rub rum 
var. hispidulum Jancz. in Monogr. d. Groseill. (1907) 2 90. — Ic. : 
A. Pojark., I.e., f. 43; Fl. Yugo-Vost. V, f. 417. - Exs. : HFR, No. 3142. 
Vernacular name: kislitsa. 

Shrubs to 2 m high, with pale shoots usually beset with stalked glands, 
often, in addition, with hairy shoots; leaves broad, with truncate or shallowly 
cordate base, dull, glabrous or sparsely hairy above, mostly pubescent 
below (f. villosum A. Pojark. ), less often glabrous (f. g 1 a b r u m 
A. Pojark.) or only with glandular pubescence (f. glandulosum A. Pojark), 
3-, rarely 5 -lobed, with broad, usually obtuse lobes, coarsely and obtusely 
dentate; petioles mostly glandular -hispid and, in addition, sometimes 
pubescent; racemes initially obliquely ascending, later nodding, 3—7 cm 
long, rather dense, 6—12 (l6)-flowered; axis and pedicels, slightly glandular, 
varying between 2.5 and 5 (7) mm long; flowers small, yellowish-greenish; 
berry 8—10 mm in diameter. Fl. end of May —beginning of June, fr. July. 

Most, mainly coniferous forests, forest edges, deciduous forest strips 
beside streams and rivers, shrub thickets, bog margins.— Arctic: Arc. 
Eur. (E.); European part: Dv.-Pech. (E.), V.-Kama. (E.), V.-Don., Transv.; 
W.Siberia: Ob, U. Tob., Irt., Alt.; E.Siberia: westernmost part of Ang. -Say.; 
Centr. Asia: Balkh., Dzu. -Tarb. Gen.distr.: Mong. (Tanno -Ola, Mongolian 
Altai). Described from Lake Bolshoe Chebache, in Kazakhstan. Type in 
Leningrad. 

Economic importance. As for preceding species. 

Note. The Yenisei River is the eastern border for R.hispidulu m; 
the western border runs to the west of the Volga, but it is impossible to 
ascertain owing to lack of data and to the difficulty in separating 
R. hispidulum from R.pubescens in that area, since they are linked 
by forms with transitional characters, apparently of hybridogenic origin. 

6. R.pubescens Hedl. in Bot. Notis. (1901) 100; Poyarkova in Tr. Bot. 
Inst. Akad. Nauk SSSR, ser. 1, No. 2 (1936) 174.- R. rub rum J3 sylvestre 
Wimm. et Grab., Fl. siles. (1827) 209.— R. rub rum j3 pubescens Swartz, 
Summa Veg. Scand. (1844) (nom. nud.). — Gro s s ula r ia rubra Rupr., 
Fl. ingrica (i860) 419.— R. schlechtendalii Lge., Ind. sem. hort. Havn. 
(1870)31 (ex parte).- R. rub rum (ex parte) Shmal'g., Fl. I, 358. - 
R. r ubr um Jancz. inComptes rendus Ac. Paris (l 900)589 (non L.). — R. c au - 
casicum R. Regel in Tr. Bot. Sada Yur'evsk Univ. II (1901) 72. (non M. B.) — 
R.warscewiczii Jancz. in Tr .Troitsko-Savsko-Kyakht.Otd.R. G. 0.(1 902 ) 
10 (non in Frut. Vilm.). — R.rubrum var. scandicum Jancz. in Monogr. 
de Groseill. (1907). — R.rubrum var. pubescens Jancz., 1. c. (ex parte). — 
Exs.: HFR, No. 1615.- Ic: Poyarkova, 1. c. (p. 1 73). 

Shrub 1—1.5 m high, with pale, initially pubescent, later glabrescent 
shoots; leaves 3- or 5 -lobed, with broadly triangular, short -acuminate, 
subobtuse lobes, glabrous or sparsely hairy above, more or less densely 
pubescent below; racemes loose, 4—9 (ll ) cm long, 8— 22 -flowered, initially 
slightly ascending, later horizontal or nodding; pedicels 3—7 mm long, more 
or less pubescent; flowers small; sepals sharply ciliate -margined, green, 
usually with copper-red or brown spots; petals of the same color; berry 
small, 6—8 (10) mm in diameter. Fl. June, fr. July. (Plate XIII, Figure 4). 



184 



Forests, mainly coniferous, forest edges, riverbanks, among shrubs. 
European part: Kar.-Lap. (?), Dv.-Pech. (W.), Lad.-Ilm., U. V., V. -Kama, 
U. Dnp., M. Dnp., V. -Don, L. Don. Gen. distr. : Scand. (S. and Centr. parts 
of Scand., Finland, Baltic States, NE Prussia), Centr. Eur. (Poland). 
Described from Scandinavia. Type in Stockholm. 

Economic importance. A whole series of red currant varieties 
originated from the crossing of this species with R. vulgare Lam. and 
R. petraeum Wulf. The fruit of the wild -growing species have a more 
acid taste than those of the cultivated currant. 

Note. A currant specimen collected in the Don, floodplain, opposite 
Belogor'e, is distinguished by the continuous whitish -tomentose pubescence 
of shoots, inflorescence axis, and underside of leaf, rather dense pubescence 
of upper leaf surface, rigid consistency and smaller size of leaves, and small 
(5—6 mm) berries. It has been isolated as a separate form — f. cretacea 
A. Pojark. nov. 

*R. scandicum Hedl. in Bot. Notis. (l 901 ) 99. — R. rubrum var. 
pubescens Jancz., Monogr. d. Groseill. (1907) 289 (ex parte). — 
R. rubrum L.Xpubescens Hedl., 1. c. 

Shrub ca. 1.5 m high; shoots pale, slightly pubescent, often in addition, 
beset with glandular hairs; leaves 3- or 5-lobed, with well developed 
coarsely dentate, acute lobes, glabrous above, usually rather sparsely 
pubescent below, with cordate or truncate base and with glandular petioles; 
racemes 2.5—5 cm long, erect or arched-upcurved, nodding in fruit, 
5—12 (l6)-flowered; pedicels 3— 8 mm long, glabrous and slightly glandular 
like the axis; flowers of medium size, green or brown; with sepals 
glabrous on the margin; berry ca. 8 mm in diameter. Fl. June, 
fr. July— August. 

Forests, forest edges, riverbanks. — European part : Kar.-Lap. and 
W. part of Dv.-Pch. Gen. distr. : Scand. (Scandinavian Peninsula and 
Finland). Described from S. Sweden. Type in Stockholm. 

Note. This name has been used to describe specimens of currants 
which, according to their morphological characters, occupy an intermediate 
place between R. r ub r um L. and R. pubescens Hedl. and are apparently 
their hybrids since their distribution area basically occupies a zone 
intermediate between the distribution areas of these 2 species, although 
partially overlapping them. 



Series 4. Meyerianae A. Pojark.— Flowers short -campanulate or 
turbinate, the hypanthium smooth inside, the sepals erect, densely ciliate, 
the style cylindrical, the ovary inferior with flat apex; berries black. 
Other species of this genus are distributed in the mountains of China, 
E. and S. Mong., and the NW Himalayas. 

7. R. meyeri Maxim, in Mel. biol. IX (1873) 232 (ex parte). - R. r ub r um 
var.foliis basi truncatis Kar. et Kir. in Bull. Soc. Nat. Mosc. XV 
(1842) 356.— R.triste Kar. et Kir., 1. c. (non Pall.). — R.atropurpureum 
Kar. et Kir., 1. c; Trautv. in Bull. Soc. Nat. Mosc. XXXIX, 1 (1866) 365 
(non C. A. M.). — Grossularia atropurpurea Osten-Saken et Rupr. 
in Mem. Ac. St. Petersb. VII se'r., XIV (1869) 47. — R. rub rum var. 



185 



Lnte r tried i u m Rgl. e1 Schmalh, m A. H. P, V (1877) 584. - K. mey eri 
vai'.iurkestauinim Jancz. in Monogr. d. Groseill. (1907) 299. — 
i\. p e i r a e u m var, a t r o p u r p u r e u m Fedtsch., Consp. El. turk. Ill ( i v>09) 
74. R. petraeum var. lit win o w i i Fedtsch., I.e. (non Jancz.). — 
Ic: Poyarkova In rr.Prikl. Bot., Gen. i Sel.XXII, 3 (1929) f.44. 
Vernacular name: chernaya kisli 

Shrub to L.5m high, with yellow, glabrous, glandular or slightly pubescenl 
shoots: loaves on fruiting shoots 2.5—5(5.5)0111 in diameter, those on 
vegetative shoo is to 6 (7) cm, orbicular, with cordate or t runcate base, 
5-lobed, loss often trilobate, with little developed, subobtuse or short- 
acuminate Lobes, glabrous on both sides (f. glabrum \. Pojark.) or 
densely pubesoent below (f. h i r t urn A. Pojark.). or glandular -hispid 
on both sides, with glandular shoots and petioles (f. g 1 a n d u 1 o S um 
\. Pojark. )j racemes short, 2 — l(5)em long, dense, 4 L2-floweredj pedicels 
1 .;"- 2 mni long, hairy; flowers turbinate, brownish with purple spots 
and nerves or dark dingy purple; berry violet -black, 7— ti mm in diameter, 
aeid. Fl. Juno, fr. August. (Plate Mil, Figure 7). 

Mountain slopes and gorges in the middle mountain zone, among shrubs, some- 
times penetrat ing into t lie subalpine .-.one. to 3,500 m, — W. Siberia : All. (SWpart, 
near lake Ma rka -kuP ): CetUr. Asia: Dzu - larb.. Syr D., Pam. -Al., T. Sh. 
Gen. distr.: Dzu.-Kash. (W.part of Kunlun, Kuldja and E.Altai near Barkul'). 
Described from the Trans-Ili Ala-Tau. Type in Leningrad. 

Economic importance. Distinguished by its abundant fruiting and pleasant 
taste of fruit. Deserves attention as a baccate plant; the fruits may be 
used like those of the cultivated red eurrant R.vulgare Lam. Ornamental 
,.„ plants. 

Note. Erroneously mentioned for the Altai in Krylov's Fl. of W. Siberia. 
The specimen from the Ulegumen River cited by Gunge belong to 
R.altissimum Turcz.; R. a t r o p u r p u r e u m var. ^ C. A. M. and 
R. petraeum var. litwinowii Jancz., reported as synonyms, also 
belong to this species. 



Series 5. Petraeae A. Pojark.— Hypanthium campanulate, smooth inside 
or with papilliform excrescences below petals: sepals densely ciliate- 
margined; style conical; ovary half-inferior with conical apex; berry 
red, dark red, or purple -black, very acid. Apart from our species, 
R. p e t r a e u m Wulf. (Alps, Carpathians, mountains of N. Africa) also belongs 
to this series. 

*8. R. latifolium Jancz. in Bull. lead. Cracov. (1906) 4 (ex parte): ibid, 
(l 913) 723 (sensu strictoi). — R. petraeum |B tomentosum Maxim, in 
Mel. biol. IX (1873) 231 (ex parte). — R.petraeum a t y pi c um Mat sum., 
Ind.pl. jap. 11,2 (1912) 187.- Ic. : Jancz. in Bull. Ac. Crac. (1913) 723, f. 5. 

Shrub 1— 2 m high; bark of shoots light brown, strongly ramentaceous; 
leaves thin, broad, with cordate base, to 12 cm long and 15 cm broad, 
3— 5-lobed, with short, broadly triangular lobes, sparsely hairy above, 
densely pubescent below, or quite glabrous on both sides, acutely dentate; 
racemes 4—8 cm long, loose, 10— 20 -flowered; pedicels 1.5—3 mm long; 
pedicels and axis more or less pubescent or glabrous; flowers 5— 7 mm long, 



ISo 



/ dark purplej sepals slightly spatulate, spreading only at the apex; 
style narrowly conical; berry red. Fl. May, fr. June. 

Undergrowth of mountain forests. — Not yet found in the Soviet Union. 
Gen. distr.: Jap. -Ch. (S. part of Sakh., Japan, Hokkaido, Honshu; S. Kurile 
Islands). Described from Honshu. Type in Cracow. 

Economic importance. Ornamental fruit plants; fruits can be used 
for the making of jam, syrup, jelly, fruit drinks, etc. 

9. R. pallidiflorum A. Pojark. in Acta Inst. Bot. Ac. Sc. URSS, ser. 1, 2 
(1936) 178.— R. at r o pu r pu r eum var.tomentosum Maxim., Prim. 

fl. Amur. (1859) 118j Fr. Schmidt, Fl. amg.-bur. II (1874) 49.- R.petraeum 
atypicum Maxim, in Mel. biol. IX (l 873) 231 (ex parte quoad pi. e. Li - 
Fudin). — R.petraeum |3 tomentosum Maxim., 1. c. (ex parte) Jancz. in 
Bull. Ac.Crac. (1913) 721.- R. lati folium Jancz. in Bull. Ac. Cracov. 
(1906)4 (ex parte). - R.petraeum Kom., Fl. Mansch. II (l 903) 440 (non 
Wulf). — R. 1 a t i f o 1 i u m Kom. and Alis., Opred. rast. Dal'nevost. kraya I 
(1931) 622.- Ic: Kom. and Alis., I.e., Plate 146. 

Shrub 1—2 rn high; pubescent or glandular -hispid shoots covered with 
strongly ramentaceous light brown bark; leaves to 10 cm broad and 9 cm 
long, thin, usually 5 -lobed, with acute or acuminate triangular lobes, 
acutely duplicato -dentate, usually sparsely hairy above, more or less 
densely pubescent to tomentose below, or else covered — sparsely above, 
densely along the veins below — with long-stalked glands, but then petioles 
and shoots also glandular-hispid; racemes 4—10 cm long, 10— 27 -flowered, 
with slender, pubescent axes and pedicels; flowers 4.5— 5 mm long, pale, 
yellowish or pinkish, with erect sepals spreading only at the apex, and 
with narrowly conical styles; berry light red, acid, ca. 8 mm in diameter. 
Fl. May — June, fr. July. (Plate XIV, Figure 3). Undergrowth, coniferous 
forest edges, in rock streams among forests; solitary or in small groups. — 
Far East: Ze. -Bu., Uda, Uss., Sakh., S. Kamch. Gen. distr. : Jap.-Ch. 
(N.Korea). Described from the Ussuri area (between the upper reaches 
of the Lifudzin and Dadansy rivers). Type in Leningrad. 

Economic importance. Ornamental plants. The berries can be used 
for the making of syrup, jam, jelly, and various drinks. 

Note. Until now, R. pallidiflorum has been identified with the 
Japanese R.latifolium Jancz.; the latter, however, is clearly distinguished 
by its dark, dingy purple flowers and by several other characters. 

R. pallidiflorum shows great polymorphism in a series of characters : 
(l ) the pubescence, varying greatly in density and character between different 
specimens — this, incidentally, is more or less peculiar to other species of 
the subgenus Ribesia; (2) size of leaves and shape of their lobes; (3) flower 
color, which is always pale but varies from pale yellow to pink. Attempts 
to ascertain the geographical character of these variations have failed, but 
it should be noted that in Kamchatka and N. Sakhalin, as compared with 
the mainland, the prevalence of specimens with very large, usually 
broad-lobed leaves is characteristic, and that specimens with leaves 
almost devoid of pubescence (not yet known on the mainland) are of 
frequent occurrence. 

10. R. altissimum Turcz. (in schedis) ex A. Pojark. in Acta Inst. Bot. 
Ac. Sc. URSS, ser. 1,2 (1936) 179; Ldb., Fl. Ross. II, 1, 198 (pro synon. 
R.petraei). - R. triste Pall, in Nova Acta Acad. Petrop.X (1797) 378 



187 



! ' 



(ex parte: pl.fruct.). R. atr opurpur euro var. J C, \. i\i. m Ldb., 
i-M. \h. [(18 >] R.altaicum Lodd. ex Baxt. in Loud.Hort.brit., 

Suppl, 11 (i 830) 667 (nom, nudum), R. triste rurci . Ln Bull. Soc. Nat. 
Mosc.X (1837)58 (non rail.): Fl.baic.-dah. I (1844) 444; tledlund in Hot. 
Not is. (1901 ) 104, R. petraeum Ldb., Fl. Ross. [1,1, 198, R.pel ra e u m 
o atropurpureuni Krylow, Fl. Ut. II (1901) 166, R. triste (ex parte) 
Kom., Fl. mansh. (1903) 142, R, petraeum var. lit wino w ii Jancz. in 
Monogr > : Groseill. (1907) 2 94. R.piMi'aoum var. a It iss imum Jancz., 
I. c. (non in Bull. \c. Cracov. L913). R. a1 ropurpureum (ex parte) Kryl., 
Fl. Zap. Sib. VI, 1441, [c: I 'oyarko'N a, 1. c, fig. 8 (p. 181), Vernacular 
name : chernaya kislil sa. 

Shrub 2 3m high; shoots glabrous or glandular -hispid, the bark dark, 
reddish brown, splitting longitudinally and separating in [ong Lamellate 
strips; leaves firm, dark green and lustrous above, lighl and whitish below, 
glabrous on both sides or else glabrous along the veins below and 
uninterruptedly glandular -hispid above, usually trilobate with slightly 

developed broadly triangular lobes, shallowlv cordate at base: petioles 
usually reddish: racemes 2.5 6(8)cm ion;',. 7 25-flowered; pedicels 

inni Long, pubescent; flowers small,! I.Smni broad, yellowish with 

dingj purple spots; sepals recurved; style broadly conical; berry purple 
black, 5 7 mm in diameter. Fl. June, fr. August. (Plate XIV, Figure 2). 

I'aluses composed Of Large stones, in the forest .one. often penetrating 

Into the bald -mountain zone, — W. Siberia; Alt.; E.Siberia: s\v Lena-Kol., 
(NW end of 1 eke Baikal), Ang -Say. and \\ . ban. Gen. distr. : Mong. (Mong. 
and ruva). Described from the Chikoi River in Dauria, rype In Leningrad. 
Note. \ species with a very complex synonymy, having been confused 

with a number of other Siberian speeies: with K.t riste Pall, which has 
red fruits and flat flowers, and with the Central \siau black-fruit ed 
K.meyeri Maxim. tin Fl. .'.ap. Sib. . t lu glandular form R. alt i S S imum 
is recorded under this name"). Botanists have not distinguished it sufficiently 

precisely from R.a1 ropurpureum C.A, M., which has red berries, 
purple flowers, different leaf shape, light bark, a different distribution area 
and different ecology. 

Economic importance. Fruits small, with thick skin, but with a pleasant 
acid taste. Of interest to fruit-growers and breeders owing to the color 
and abundance of its fruits. 

11. R. atropurpureum C. A. M, Ln 1 db„ I'd. Alt. 1 (1829) 268 (ex parte: 
var, q e1 •) ); \. Pojark, in Bull, of Appl. Bot., Gen. et Plant -Breed. XXII, 3 
(1929) 346, - R. triste (non Pall.) Bge., Suppl. Fl. altaic. (1835) 21.- 
R. triste var, r foliis subtus puberulis Bong, et Meyer in Mem. Ac. Sc. 
St. -Petersb. Se*r. VI. IV (1845)191.- R.baicalense Turcz. in schedis 
(pro parte), - R. p e t ra e u m (ex parte) Ldb., Fl. Ross. II, 1, 198. — 
R. a t r o p u r p u r e u m var, t y p i c a Trautv., Enum. pi. song. 11 (l 866 ) 40. — 
R. petraeum a typicum Maxim, in Mel. biol. l\ (1 873 ) 230 (ex parte). — 
R. petraeum o rub rum Kryl., Fl. Alt. II (l 901 ) 465. - R. p e t r a e u m 6 
a t r p U r p u r e u m Jane.-, in Monogr. d. Groseill. (1 907) 293. — R.atro- 
145 purpureum var. rubrum Kryl., Fl. Zap. Sib. VI (1931 ) 1442. — Ic. : Ldb., 
[C. pi. Fl. Ross. tab. 231 : Foyarkova. I.e., Fig. 46. 



ISS 



246 



Shrub 1—1.5 m high, with light, grayish yellow shoots; leaves rounded- 
cordate, 8—10 cm in diameter, thin, glabrous on both sit ow, 
3 or 5-lobed, with acute, well developed upper lobes and inconspicuous 
lower Lobes, often acutely and coarsely duplicato-dentate; racemes 
2 4.5(6)cm long, mostly dense, 4—15 (20)-flowered; flowers 4— 5mm Lc 
purple, rarely Light writh purple nerves; style broadly coi Leal; berry 
red, '<'> L0 mm in diameter, sometimes to 11— 13 mm. Fl. May — June, 
fr. July* August. 

Shady slopes and banks of streams and rivers, in mountains and in the 
Lowland forest zone.- W. Siberia: Ob (S. part), Irt. (SE), Alt.; E. Siberia: 
STeniS. (S. part), Ang. -Say.; Centr. Asia: Dzu.-Tarb. (Tarbagatai, Saurj. 
Gen. distr.: Mong. (Mongolian Altai, Tannu-Ola Range). Described from 
the Ursul River in the Altai. Type in Leningrad. 

Hybrids: R. a t r op u r p u r e u m C. A. M. X a It i s s i m um Turcz., 
apparently rare. The leaves are firm, bicolor as in the latter species 
but with acute lobes and with pubescence below; bark very light brown, 
lighter than in R. altissimum and not strongly peeling; berries black 
hi herbarium specimen. In Altai and on the Slyudyanka River near Lake 
Baikal. 

Economic importance. May be of importance as a baccate plant. 

12. R. biebersteinii Berl. in Me'm. Soc. phys. hist. nat. Geneve 111,2 (1826) 
60 (nom. nudum); DC., Prodr. Ill (1828) 482; Grossg., Fl. Kavk. II (1930) 
242.- R. cau casicum M. B.,F1. taur.-cauc. Ill (1819) 160 (non Adams). - 
R. rub rum C. A. M., Enum. pi. cauc.-casp. (1831) 154 (non L.).— 

R. cilia turn C.Koch in Linnaea XVI (1842) 355.- R. petraeum Ldb., 
I I. Koss. II, 1, 198 (ex parte); Boiss , Fl.Or. II (1872) 816 (non Wulf); 
Medvedev, Der. i kust. Kavk. (1919) 166. — R. petraeum var. caucasicum 
Jancz. in Monogr. d. Groseill. (1907) 293.- Exs.: PI. Or. Exs., No. 186. 

Shrub to 2 m high with light, glabrous shoots; leaves thin, deeply cordate, 
large (to 13 cm broad and 10cm long), usually 5-lobed, either glabrous on 
both sides (f. glabr um Grossh., i.e.), or densely hairy below (f.hirtum 
Grossh., 1. c), rarely with scattered glandular bristles above and with hairs along 
the veins below and on the petioles; racemes 4—12 cm long,horizontal, nodding in 
fruit, loose, 15— 50 -flowered; pedicels 2— 3 mm long; flowers 5— 6 mm long, 
dark purple; sepals recurved; hypanthium with conspicuous excrescences 
below petals; styles broadly conical; berry small, 6—7 (8) mm in diameter, 
dark red (mature ?) or black -purple. Fl. June, fr. from end of August. 

Subgenus 2. HERITIERA Jancz. (pro sect.) in Bull. Acad. Cracov. (1906) 7 
et in Monogr. d. Groseill. in Me'm. Soc. phys., hist. nat. Geneve XXXV (1907) 
246. — Flowers in racemes, bisexual, flat; stamens attached below petals; 
style very short, deeply split. Prostrate, unarmed shrubs. Bud scales soft, 
herbaceous; glands stalked crystalline. Berry glandular -hispid, red or 
black-red. Apart from the USSR species, there are 8 more species in 
the mountains of western North America. 

13. R. sachalinense Nakai in Bot. Mag. Tokyo, XXX (1916) 144; Kom. and 
Alis., Opred.rast. Dal'nevost. kraya I (1931 ) 621; Poyarkova in Tr. Bot. 
Inst. Akad. Nauk SSSR, I ser., 2 (1936) 199.- R.alf ine var. sachalinense 



189 



247 



Fr. Schmidt, Fl. sachal. (1874) 144. — R. lax if lo rum (non Pursh) Maxim, in 
Mel. biol. IX (1873) 227; Jancz. (ex parte) in Monogr. d. Groseill. (1907) 
306. 

Small shrub with decumbent and rooting stems; leaves 4—9.5 cm broad, 
orbicular, deeply cordate, glabrous, with long-stalked glands — which cover 
also the petioles, inflorescence axis, and pedicels — only along the veins 
below; lobes 5— 7, large, oblong-rhomboid, acute, duplicato-dentate; racemes 
to 10 cm long, 5—1 5 -flowered; flowers purple, with recurved sepals; berry 
red, glandular -hispid. Fl. June, fr. July. (Plate XV, Figure 1 ). 

Moist forests and riparian shrub thickets.— Far East: Sakh. Gen. distr.: 
Jap.-Ch. (japan). Described from the island of Shikoku, Japan. Type in 
Tokyo. 

Economic importance. Fruit edible, rather acid, pleasant to the taste. 
Deserves to be cultivated. 

14. R. malvifolium A. Pojark. in Bull. Appl. Bot., Genet, and Plant. Breed. 
XXII, 3 (1929) 352, f. 52. 

Very low shrub; leaves rounded -cordate or reniform, hairy or glandular - 
hispid above, with numerous stalked glands and scattered viscous sessile 
glands along the veins below, shallowly 5 -lobed, with rounded, obtusely 
duplicato-dentate lobes; racemes 5— 10 -flowered; flowers pinkish; sepals 
coarsely glandular on the margin; pedicels long, covered — like the axis — 
with stalked glands. Fruit unknown. End of flowering: 12 August. 

Stony taluses, at 1,200 m. Centr.Asia: Pam.-Al. Only one habitat is 
known, on the northern slope of the Gissar Range, in the upper reaches 
of the Iskander -Darya River (Komarov, 1 892). Type in Leningrad. 



Subgenus 3. GROSSULARIOIDES Jancz. in Bull. Acad. Cracov. (l 903) 238. - 
Flowers white or pinkish, in racemes, bisexual, with flat, broad hypanthium 
Berries glandular -hispid. Bud scales membranous; glands stalked, 
crystalline. Shoots spiny; spines numerous. Three species in all, of 
which two — R.lacustre Poir and R.montigenum Mc. Clatch — in 
N. America. 

15. R. horrid um Rupr. in Maxim., Primit.fl.amur. (1859) 117; Kom., Fl. 
Mansh. II (1903) 446; Kom. and Alis., Opred. rast. Dal'nevost. kraya I (1931) 
622.— R.lacustre var. horridum Jancz. in Bull. Acad. Cracov. (l 903) 
238.- Ic: Nakai, Fl. sylv. koreana XV (1926) tab.X.- Exs.: Herb. Fl. URSS, 
No. 3143. 

Very spiny shrub, 1— 1.5 m high; spines up to 20, often arranged in 2 or 3 
layers; leaves rounded -cordate, 5-, rarely 3 -lobed, bluntly duplicato- 
dentate, with acicular spines covering them above; racemes to 4 cm long, 
4— 6- flowered, the axis and pedicels covered with red stalked glands; 
flowers with broad, recurved sepals and flabellate petals; hypanthium 
purple; ovary densely glandular; berry black, glandular-hispid. FL June, 
fr. July. (Plate XV, Figure 2). 

Mountain slopes. — Far East: Uss. (coast), Sakh. Gen. distr. : Jap.-Ch. 
(N.Korea and S. Sakh.). Described from Khadzhi-bau (.now Sovetskaya 
Gavan'). Type in Leningrad. 



190 



Economic importance. Berries edible, succulent, with acid taste. In 
view of its original habitus, this shrub can be used for horticultural purposes, 
purposes. 



Subgenus 4. EUCOREOSMA Jancz (pro sect.) in Bull. Acad. Cracov. (1906) 
7; Berger in New York St. Agric. Exper. Stat. Techn. Bull., No. 109 (1929) 
29.— Flowers bisexual, in racemes, with cup-shaped or campanulate or 
goblet -shaped hypanthium and half -inferior or inferior ovary. Bud scales 
soft, herbaceous. Glands punctate, yellow, aromatic. 



Series 1. Nigrae A. Pojark. (Black currants).— Flowers campanulate or 
cyathiform, with oval petals. Leaves large, cordate, with well developed 
lobes. Berry black, aromatic. Shrubs 0.75— 1 .5 m high. Series distribution 
area: from France and Great Britain to the Pacific Ocean; apparently 
only the above-mentioned species. 

Economic importance. Berries of all species edible, differing little in 
taste from the cultivated black currant; R. nigrum L. is the ancestor 
of numerous strains and its Siberian variety the ancestor of others. 
May be used for the same purposes as R.nigrum. 

*16. R. ussuriense Jancz. in Bull. Acad. Cracov. (l 906) 12; Poyarkova 
in Tr.Bot. Inst. Akad. Nauk SSSR I, ser.2 (1936) 185.- Monogr. d. Groseill. 
(1907) 349.— Ic: Jancz., Monogr. f. 79, 80; Poyarkova, 1. c, f. 9. A. 

Branching shrub ca. 1 m high, producing abundant root suckers, with 
golden -yellow hairy shoots; leaves to 10 cm broad, firm, bright green, 
lustrous, initially scattered -hairy above, later glabrous, pubescent below 
along the veins, acutely dentate, 5 -lobed, with acute or acuminate lobes; 
racemes to 2.5 cm long, 5— 9 -flowered; pedicels 3—5 mm long, hairy; 
bracts lanceolate, 2— 2.5 mm long; flowers pale or yellowish, 7— 8 mm 
long, glandular and not densely hairy outside, with slightly pentagonal 
hypanthium 2—2.5 times as broad as high; style entire; ovary broadly 
conical at the apex, densely glandular in lower part; berry black, small, 
with slender, nodding stalk. Fl. June, fr. July. 

This species has not yet been found in the Soviet Union. Only one habitat 
is known, viz., on the Segelsu River, an affluent of the Tumen in N. Korea. 
Described after a cultivated specimen. Type in Cracow. 

Economic importance. Baccate plants with small, irregularly maturing, 
soon deciduous fruits, differing little in taste from other species of this 
series. 

Note. R. ussuriense has been described after a cultivated specimen 
grown in the Cracow Botanical Garden from seeds received from Khabarovsk 
and allegedly gathered in various unknown sites in the Ussuria region. 
The black currant growing in the vicinity of Khabarovsk and in the Amur 
Valley is identical to the Daurian R.pauciflorum Turcz., and differs from 
R. ussuriense Jancz. in a series of characters enumerated in the key. 
There is only one specimen from Lake Khanka in the Ussuri region; it has 
been collected in a sterile state and therefore cannot be determined. 



191 



(249) 




PLATE XIV. 1 — Ribes ma nshu ri c u m (Maxim.) Kom., flower; 2 — R. a lti ssi mu m Turcz., 

a) flower, b) flower section; 3 — R.pallidiflorum A.Pojark., flower; 4— R.nigrum L., a) flower; 

5 - R. dikuscha Fisch., a) flower; 6 — R. procumbens Pall., a) flower. 



192 



251 



17. R.pauciflorum Turcz. ex Pojark. in Sched. Herb. Fl. URSS, X (1936) 
69; Ldb., Fl. Ross. II, 1, 200 (pro synon. R. nigri). — R. nigrum Turcz., 
Fl. baic.-dah. 1,1844,445 (non L.); Kom., Fl. Mansh. II (l 903) 435. - 

R. n i g r u m var. praecox E. Wolf in Mitt, deutsch. dendr. Gesellsch. 
XXXIV (1924) 331.— R.ussuriense (non Jancz.) Kom. and Alis., Mai. 
opred. rast. D. Vost. (1925) 250; Opred. rast. Dal'nevost. kraya I (1931) 621. - 
Ic: Poyarkova in Tr.Bot. inst. Akad.Nauk SSSR I, ser.2 (l 936), Fig. 9. 

Shrub to 1 .5 m high, with yellowish gray, pubescent shoots with numerous 
root suckers; leaves 5—8 (9) cm in diameter, glabrous above, pubescent 
along the veins below, 5 -lobed, the lobes short, broadly triangular, subobtuse, 
the middle lobe usually larger than the lateral; racemes 1.5—3 cm long, 
(2)3—6 (8)-flowered; pedicels 1.5—3 (4)mm long, densely pilose like the 
axis; bracts linear -lanceolate or lanceolate, 2—3 (4) mm long; flowers 
yellowish, small, 6—7 (8) mm long, densely pubescent outside; hypanthium 
short, hemispherical, smooth, twice as broad as high; sepals ligulate, 
2—2.5 times as long as broad, acute or obtuse, densely hairy outside, 
sometimes slightly flesh-colored; style about as long as stamens, entire; 
ovary high -conical, its lower part densely glandular; berry large, 
ca. 10— 13 mm in diameter. Fl. May— June; fr.July. 

Mixed and deciduous mountain and floodplain forests and forest edges; 
solitary or in small thickets; often settles on dry forest burns, where it 
grows abundantly. — E. Siberia: Lena-Kol. (S.part), Eastern part of Ang.-Say. 
Dau. ; Far East: Ze. -Bu., Uda, Uss. ? Gen. distr. : Manchuria. Described 
from Dauria. Type in Leningrad. 

Economic importance. Deserves attention as an excellent baccate plant 
with large tasty, abundant fruit and a great capacity for vegetative 
proliferation by means of root suckers. 

18. R. kolymense Kom. ex Pojark. in Acta Inst. Bot. Ac. Sc. ser., I, 2 
(1936)111; Kom., Vved.vo Fl. Yakut. I (1926) 142 (nom. nud.). - R. n ig r um 
var. koly me ns e Trautv. Fl. ripar. kolym. (1878) 34. — Ic. : Poyarkova, 
I.e., Fig. 10 (p. 189). 

Shrub; young shoots pubescent, yellow, the annual shoots subglabrous, 
brownish; leaves dull, hairy along the veins on both sides, to 6.5 mm broad 
on flowering shoots, trilobate, with long, acute, coarsely dentate lobes, 
rounded or slightly cordate at base; leaves on sterile shoots to 10 cm 
broad, obtusely lobed, with large, obtuse, mostly simple teeth on the 
margin; racemes short, 2— 3.5 cm long, 3— 5 -flowered; pedicels (2.5)3— 5 mm 
long, hairy like the axis; flowers 9—10 mm long, cyathiform, densely 
pubescent outside and, in addition, glandular; hypathium 1.5—2 times as 
high as broad; sepals erect, their upper part slightly recurved, narrow 
(2.3—3 times as long as broad); styles split to the middle; ovary high- 
conical, with glandular lower part. July. 

Rocky slopes.— E. Siberia: Lena-Kol. Only one habitat on the banks of 
the Kolyma River (above Sredne-Kolymsk) is known. Type in Leningrad. 

19. R. turbinatum A. Pojark. in Bull. Appl. Bot., Genet, a. Plant. Breed. 
XXII, 3 (1929) 34 9. — R. n igr um var.pauc if lor um Jancz. (ex parte) in 
Monogr. (1907) 341.— Ic: Poyarkova, 1. c, Fig. 50. 

Shrub; shoots pubescent, initially brownish yellow, later nearly gray; 
leaves to 10 cm broad, firm, glabrous, lustrous, acutely lobed and acutely 



193 



253 



dentate; racemes ca. 3 cm long, 4 — 7 -flowered, with glabrous axis and 
pedicels; bracts lanceolate, 2— 3 mm long; flowers ca. 9 mm long, flesh - 
colored, with scattered pubescence and slightly glandular outside; 
hypanthium pentagonal, tapering below the sepals, constricted (turbinate) 
at the apex, about as high as broad; sepals broad, only 1.5 times longer 
than broad, obtuse, sometimes overlapping; style glabrous. June. 

Riparian shrub thickets. — W. Siberia: Alt.; Centr.Asia: Dzu.-Tarb. 
Known only from the Narym Range and the Dzungarian Ala-Tau. Described 
from the Narym Range. Type in Leningrad. 

Note. Species requiring further study and observation in situ; differs 
sharply from the Siberian black currant in the shape of its hypanthium and 
its broad sepals. 

20. R. janczewskii A. Pojark. in Bull. Appl. Bot., Genet, and Plant. Breed. 
XXII, 3 (1929) 346, f.47. 

Shoots glabrous or scattered -hairy, golden, later dingy yellow; leaves 
deeply cordate, thin, lustrous, glabrous, to 15 cm broad, 5-lobed, with large, 
acute or acuminate lobes, acutely dentate, often duplicato -dentate or 
incised -dentate; racemes to 5 cm long, 5— 10 -flowered, with glabrous axis 
and pedicels and with lanceolate bracts; flowers to 12 mm long, 
flesh-colored, pubescent and glandular outside; hypanthium campanulate, 
about as high as broad; sepals obtuse, 2—2.25 times as long as broad; 
petals broadly oval, almost without claw; berry large, to 13 mm in 
diameter. Fl. June, fr. August. 

Gorges and mountain river valleys, to 3,000 m.— Centr.Asia: Pam.-Al. 
(Zeravshan, Gissar, Vakhan, and Shugnan ranges), T. Sh. (W. Tien Shan and 
Kirghiz Ala-Tau (former Aleksandrovskii Range)). Endemic. Described 
from Kul'-i-kalon in the Zeravshan Range). Type in Leningrad. 

Economic importance. Excellent baccate plants with large, aromatic 
fruits. Ornamental owing to large, lustrous, deeply lobed leaves. 

21. R. nigrum L., Sp.pl. (1753) 201; Ldb., Fl. Ross. II, 1,200 (ex parte); 
Shmal'g., Fl. Sr Ross. I (1895) 358.- R.olidum Moench, Meth.pl. 
(1794) 683.- Botryocarpum nigrum Rich., Bot. Me'd. II (1823) 490. - 
Grossularia nigra Rupr., Fl. ingrica (1859) 41 8. — Ic. : Rchb., Ic. 

Fl. germ. XXIH, tab. 137. 

Shrub 1—1.25 m high; shoots pubescent, initially pale, becoming light 
brownish toward end of summer; leaves dull, glabrous above, pubescent 
along the veins below, to 10 (l2)cm broad, 3-, less often 5-lobed, the lobes 
usually broadly triangular, the middle lobe often elongated; racemes 
3—5 (8) cm long, 5— 10 -flowered; pedicels glabrous or pubescent, 3— 8 mm 
long; bracts 1—2 (3)mm long, varying from oval to linear -lanceolate; flowers 
7—9 mm long, lilac or pinkish gray, mostly densely pubescent outside; 
hypanthium hemispherical-campanulate, as broad or 1.5 times as broad 
as high; sepals recurved, subacute, rather broad (2—2.25 times longer 
than broad); style usually entire; berry ca. 10 mm in diameter, sometimes 
brown or greenish. Fl. May — June, fr. July. (Plate XIV, Figure 4). 

Riparian thickets, moist forests and forest edges, alder stands, bog 
margins, moist meadows; solitary or in small thickets.— Arctic: 



194 



Arc. Eur.; European part: Kar.-Lap., Dv.-Pech., Lad.-Ilm., U. V., V.-Kama, 
U. Dnp., M. Dnp., V.-Don, Transv.; Caucasus: Cisc. ?, S. Transc. (on the 
Arpa-chai River and its affluent the Istisu in the Daralagez — Armenia); 
W.Siberia: Ob, U. Tob., Irt., Alt.; E.Siberia: Yenis., Lena-Kol. (SW), 
Ang.-Say.; Centr.Asia: Balkh., Dzu. -Tarb. Gen. distr.: Mong. (NW). 
Described from Sweden. Type in London. 

Note 1. While the typical form has campanulate flowers, 
var. sibiricum W. Wolf (Mitt, deutsch. dendr. Ges. XXXIV (1924) 331; 
Kryl., Fl. Zap. Sib. VI (l 931 ) 1445.— R. nigrum var. pauciflorum Jancz. 
in Monogr. (1907) 347 (ex parte). — R. cyathiforme A. Pojark. in Bull. 
Appl.Bot., Genet, and Plant Breed. XXII, 3 (1929) 344, f. 51 ), is distinguished 
by its narrower flowers, the hypanthium 1.5 times or more longer than 
broad, narrower sepals, and usually split style. This variety occurs only 
in a certain part of the distribution area of R. nigrum L., namely in the 
mountain regions of Kazakhstan where it is widely distributed together with 
the typical form (var. e u ro p a e u m Jancz., 1. c). In southern Siberia it 
occurs infrequently, and it is difficult to separate the two forms, which are 
linked by transitions. 

N o t e 2 . No reliable information regarding the occurrence of R.nigrum 
in the Caucasus has been available until recently, although in 1924 E. Wolf 
(in IVIitteil.d. deutsch. dendr. Ges. XXXIV (1924) p. 330) described the variety 
R.nigrum var. c a uc a s i c um E. Wolf after a specimen grown from 
seeds allegedly received from the Caucasus. For this specimen he reports 
1—4 -flowered racemes, with pedicels shorter than flowers, hemispherical- 
campanulate hypanthium, light green, sometimes slightly pigmented sepals, 
dentate or even lobate petals, often irregular in shape, and even several 
properties concerning the color and shape of buds and flower-buds. The 
black currants I found in 1936 in the Daralagez Range (Armenia), on the 
rocky banks of the Arpa-chai and Istisu rivers, were collected during 
fruiting and therefore could not be compared fully to var. caucasicum, 
but the following differences from the latter may be noted: more abundant 
4— 9-flowered raceme and entire, acute, ligulate, short-clawed petals of 
regular shape. Thus, the question of the Caucasian mountain currants 
versus the European currants is an issue to be settled when more 
complete material is available. 

Economic importance. R.nigrum L. is the ancestor of nearly all 
varieties of cultivated black currants. It is very valuable for the abundant 
vitamin C in its berries. The berries are eaten raw and are used for making 
jam, syrup, jelly, canned food, candy, marinades, refreshing drinks, 
fruit wine, infusions, and liqueur. The leaves are used for vegetable 
preservation and sometimes as a tea substitute. Percentage chemical 
composition of berry sap according to Hegi: 36.1 — water, 10.4—12.8 sugars, 
2.7—3.7 free acids, (mainly malic acid), small quantity of pectins, 0.7—0.93 — 
ash; ash content: 1 6.4% CaO, 13.8% P 2 O s , 5.2% MgO. The leaves, and 
especially the flowers and buds, contain essential oil. 

Ornamental shrubs. Variegated- and laciniate-leaved forms are known 
(f. a p i i f o 1 i u m and f. a c o n i t i f o 1 i um Kirchn). 



Series 2. Dikuschae A. Pojark. — Flowers white, flat, with flabellate 
petals, cylindrical style, and inferior ovary. Berry black-blue or black, 



195 



not aromatic. Shrubs with large, orbicular leaves, the leaf lobes well 
developed.— The N.American R.petiolare Dougl. (mountains of the 
Southwest) and R. hudsonianum Rich. (Canada — from Alaska to 
the Hudson Bay) are closely related species. 

22. R. dikuscha Fisch. in Turcz., Fl.baic.-dah. I (1842) 445; Trautv. 
et Mey., Fl. ochot. (1856) 40; Maxim, in Mel. biol. IX (1873) 223; Kom., Fl. 
Manch. II (1903) 446; Wed. v izuch. rast. Yakut. I (l 936) 142; Kom. and 
Alis., Opred. rast. Dal'nevost. kraya I (l 931 ) 621.— R. americanum Pall. 
Fl. Ross. I, 2 (1788) 34 (non Mill.).- R.appendiculatum Kryl. in 
Sched.Herb. Fl. Ross. VI (1908) 128.- Ic: Poyarkova in Tr. Bot. Inst. 
Akad. Nauk SSSR, 1 ser. 2 (l 936), Fig. 1 1 . - Exs. : HFR, No. 1929. 
„j-c Vernacular name : aldanskii vinograd. 

Shrub; leaves to 13 cm long, 10 cm broad, light, slightly glaucous -green, 
glabrous on both sides or pubescent along the veins below, acutely 3- or 
5-lobed, with cordate base; racemes to 8 cm long, 8— 13 -flowered, loose; 
pedicels slender, 3— 7 (12) mm long, glabrous; flowers large, to 9 mm in 
diameter; sepals tomentose-pubescent on both sides; hypanthium and 
ovary glabrous outside; berry to 13 mm in diameter, blue-black with 
waxy bloom. Fl. June, fr. August. (Plate XIV, Figure 5). 

River islands, river alluvium and stony banks, riverain deciduous forest 
strips, including their edges; often in small thickets.— Arctic: An.; 
E.Siberia: Lena -KoL, Dau. (NW part); Far East (rarely): Ze.-Bu. 
(isolated sites in the upper reaches of the Zeya River, on the Selemdzha, 
Urkan, and Bureya rivers), Uda, Uss. (known only from the Botchi River). 
Endemic. Described from Upper Angara River. Type in Leningrad. 

Economic importance. The berries of this species are not aromatic 
and resemble Va c c in i um uli'gonosum L. Distinguished by very 
large fruits and abundant fruiting. The crossing of these currants with 
other species of the subgenus Eucoreosma — distinguished by the 
excellent taste of their berries — promises good results. 



Series 3. Procumbentes A.Pojark.— Flowers flat, purple, with flabellate 
or obtriangular petals, conical style, and half-inferior ovary. Berry 
brown. Low shrub. One species. 

23. R. procumbens Pall., Fl. Ross. I, 2 (1788)35; Turcz., Fl. baic. -dah. I, 
446; Ldb., Fl. Ross. E, 1, 198; Kryl., Fl. Zap. Sib. VI, 1444; Kom., Fl. 
Mansh. II (1903)436; Kom. and Alis., Opred. rast. Dal'nevost. kraya I (1931) 
621.- Ic: Pall., 1. c.,tab. 65; Poyarkova in Trud. Bot. Inst. Akad. Nauk SSSR, 
I ser. 2 (1936), Fig. 13.- Exs.: Herb. Fl. URSS, No. 3146a, 3146b, 3146c. 

Low shrub to 25 cm high; branches prostrate, often decumbent and 
rooting; lustrous golden shoots beset with sparse punctate glands; 
leaves glabrous, to 6 cm long and 8 cm broad, dark green, lustrous, and 
smooth above, light, whitish, with sparsely scattered glands below, orbicular 
or rounded -reniform, 3- or 5-lobed, with mostly rounded, rarely subacute, 
coarsely dentate lobes and with cordate or truncate base; raceme short, 
6— 10 -flowered, the axis and pedicels glabrous; bracts very short or 
absent; flowers 5— 8 mm in diameter; hypanthium greenish, glabrous outside 



196 



or with few glands; sepals purple, densely tomentose -pubescent on both 
sides; berry ca. 1 cm, sepals purple, densely tomentose -pubescent on 
both sides; berry ca. 1 cm, sometimes to 12—13 cm in diameter, greenish, 
dark brown near the tip, very aromatic. Fl. June, fr. July — August. 
(Plate XIV, Figure 6). 

Well drained sites with moistened soil, stream and spring banks, moist 
forests, stony moss-covered soils. — W. Siberia: Irt., Alt., Ob (S.); 
E.Siberia: Yenis.(S.), Lena-Kol. (S.), Ang.-Say., Dau.; Far East: Okh., 
Ze.-Bu., Uda, Uss. (very rarely; known from the Botchi River and from 
Sovetskaya Gavan), Sakh. ? Gen. distr. : Mong. (Tuva ASSR, N. Mong.) 
and Jap.-Ch. (N. Korea, Manchuria). Described from Dauria. Type in 
London. 

Economic importance. Distinguished by its exceptional gustatory 
qualities. Extensively used by local population for the making of jam, 
liqueur, etc. Sometimes cultivated by local amateurs and horticulturists. 
Fully deserves to be widely cultivated. 



Series 4. Fragrantes A. Pojark.— Flowers flat, white, with flabellate 
petals, cylindrical style, and inferior ovary. Berry brown. Low shrubs 
with reniform leaves, the lobes slightly developed. Two species in the 
USSR. 

24. R.fragrans Pall, in Nova Acta Ac.Petrop.X (1797) 377; Turcz., 
Fl. baic.-dah. 1,447; Kom., Fl. Mansh. II (1903) 437; Jancz. in Monogr. d. 
Groseill. (1907) 343 (ex parte); Kom. and Alis., Opred. rast. Dal'nevost. 
kraya I (1931) 621.— R.pneobalsamum Sievers in Neuerst. nord. Beitr. 
Ill (1796) 218 (nom. nud.). - R. f ragran s a glabr um Ldb., Fl. Ross. II, 

1 (1844) 197.— R. suaveolens Turcz., Fl. baic. -dah., 1. c. (pro synon.). — 
Ic: Pall., I.e., tab. 9; Lodd., Bot. cab. XVI (l 829) tab. 1533. - Exs. : H. Fl. 
URSS, No. 3141a, 3141b. 

Shrub not more than 50—70 cm high, with ascending branches and densely 
glandular shoots; leaves coriaceous, to 6 cm broad, dark green, glabrous, 
and lustrous above, densely covered with glands below, trilobate, with 
slightly developed, sometimes inconspicuous rounded or rounded -triangular 
lobes, acutely dentate; racemes to 5 cm long, 6— 12 -flowered; pedicels 
long, 2—4 mm, densely glandular and sometimes, in addition, hairy; sepals 
arachnoid -pubescent on both sides; hypanthium greenish, glabrous outside 
or glandular; berry red-brown (black at maturity), ca. 8 mm in diameter, 
sometimes bitter. Fl. June, fr. from end of July. 

Mountain slopes, among rocks and on stony taluses, to lower part of 
the bald -mountain zone.— E.Siberia: Lena-Kol., Ang. -Say. (E.), Dau.; 
Far East: Okh., Ze.-Bu., Uss. Gen. distr. : Mong. (NE). Described from 
the Chikoi River in Transbaikalia. Type in London. 

25. R. graveolens Bge. in Suppl. Fl. Alt. (1835) 19; Kryl., Fl. Zap. Sib. VI, 
1443.— R.fragrans Bong, et Meyer in Mem. Ac. Sc. St. Peter sb. VI, 

ser. Sc. nat. IV (1845) 191 (non Pall.). — R.fragrans /3 infracanum 
Ldb., Fl. Ross. II, 1 (1844) 1 97. - R. f r a g r a n s (ex parte) Jancz. in 
Monogr. d. Groseill. (1907) 343. 



197 



257 



258 



Low shrub with ascending branches and hairy, densely glandular shoots; 
leaves small, 1—3 (5) cm broad, firm, dull and rugose above, densely covered 
below with punctate glands in addition to tomentose pubescence; racemes 
2—4 (5) cm long, 4— 10 -flowered, the axis and pedicels glandular and densely 
pubescent; berry 8—10 mm in diameter, brown. Fl. June— July, fr. August. 

Subalpine zone, stony slopes and rock streams, also sometimes in 
gravelly and moss -and -lichen tundra.— W. Siberia: Alt.; E. Siberia: 
Ang.-Say. Gen. distr. : NW and N. Mong. Described from the Chuya 
River in the Altai. Type in Leningrad. 



Subgenus 5. BERISIA Spach, Hist. nat. d. Veg. VI (1838) 167; Jancz., 
Monogr. d. Groseill. (1907) 259.— Flowers in erect racemes with large 
bracts, dioecious, the staminate with almost wholly reduced ovary, the 
pistillate with abortive stamens; petals very small; bud scales membranous; 
glands stalked and crystalline or viscous and sessile. 

Economic importance. The berries are unpleasant to the taste, inedible 
when raw, but can be used for the preparation of fruit drinks and fruit wine. 
Some species are ornamental. 



Series 1. Orientales A.Pojark.— Flowers hairy outside, whitish or 
purple; berries sometimes hairy or slightly glandular. Low, unarmed 
shrubs with pubescence of spreading hairs, usually in addition to viscous, 
aromatic, sessile glands and crystalline stalked glands. Distribution 
area of the series: E. Med. and mountains of Middle Asia. 

26. R. orientale Desf., Hist. d. arbr. II (1790) 88; Ldb., Fl. Ross. II, 
1,197; Grossg., Fl.Kavk. II (1931) 241.- Exs.: Fl. cauc. exs., No. 136. 

Shrub to 1 m high; leaves 1.5—4.5 cm broad, usually broad, with truncate 
base, dark green, lustrous, 3- or 5-lobed, partly coarsely duplicato -dentate, 
hairy on both sides but more densely so below, with, in addition, 2 types 
of glands: sessile and viscous, scattered on both sides of leaves and 
communicating a resinous odor to the leaves, and stalked crystalline 
glands, usually densely covering, in the form of bristles, the upper 
surface of the leaves and the veins on the underside, and in the form 
of glandular hairs covering the petioles, inflorescence axis, and pedicels, 
which, in addition, are rather densely pubescent with simple spreading hairs. 
Flowers white (flower buds sometimes brownish-reddish); berries red, 
glabrous or covered with sparse glands. Fl. May, fr. July. 

Rocky sites on mountain slopes in the 1,200— 2,000m belt.— Caucasus: 
Cisc, Main Range, W., E., and S. Transc, Dag., Tal. Gen. distr. : Arm. -Kurd. 
(Turkish Armenia), Bal.-As. Min. (As. Min., Greece), E. Med. (Lebanon). 
Described from Lebanon. Type in Paris. 

Note. In his monograph, Jane zewski, who understood the species 
R. orientale Desf. sensu latb., and who groups under this name all 
species of the series Orientales, separates the Caucasian plants as 
a distinct variety, R. orientale var. genuinum Jancz., indicating that 
they are distinguished from the Lebanese plants (R. orientale var. 
resinosum Jancz.) by their dark green, lustrous, flat leaves, with mat, 
grayish, slightly rugose leaves. 



198 



259 



Due to the lack of herbarium specimens from Asia Minor, Lebanon, 
and Greece, it is impossible to verify and evaluate the taxonomic significance 
of the differences indicated nor has it been possible to ascertain whether 
in the E. Mediterranean area one is dealing with one species, or whether 
here — as in Middle Asia — the currants of the series Orientales are 
differentiated into several geographic races. 

27. R.melananthum Boiss. et Hoh. ex Kotschy, PL Pers.bor. 1846; Boiss. 
Diagn. I Ser. X (1849) 19; Pojark. in Bull. Appl. Bot. Gen. and PL Breed. XXII, 
3 (1929)339; in Acta Inst. bot. Acad. Sc. URSS, 1 ser., II (1936) 203. 

Low shrub with crowded, curved branches; leaves small, 1—2 cm 
broad, slender, pubescent on both sides, incised to the middle into 3 obtuse 
lobes, with few large teeth on the margin and with cordate base; petioles 
with pubescence of spreading hairs with an admixture of glandular hairs; 
inflorescence axis and pedicels hairy; bracts oblong-oval, glandular on the 
margin; flowers dark purple; berries unknown. 

Rocky mountain slopes; vary rare.— Centr.Asia: Mtn. Turkm. 
Gen. distr.: Iran. (Mt. Elburz). The single sterile currant specimen 
supposedly belonging to this species has been collected on Mt. Chapan-Dagh 
in E. Kopet Daghandis identical to the original specimen R.melananthum 
Boiss. et Hoh. except for the rather abundant pubescence on the petioles. 
No other sites are known in the Soviet Union. Described from the 
Naseron Range in the Demavend and Elburz mountains and not reported 
from anywhere else. Type in Geneva; cotype in Leningrad. 

28. R. heterotrichum C. A. M. in Ldb., Fl. Alt.I (1829) 270; Ldb., Fl. 
Ross. II, 1, 197; KryL, Fl. Zap. Sib. VI, 1448. - G r o s s u 1 a r i a adeno- 
p hy 1 1 a Osten-Sacken et Rupr. in Mem. Acad. St. Pe'tersb. Se'r. VII, XIV 
(186 9) 47.— R. orientale var. heterotric hum Jancz. in Monogr. d. 
Groseill. (1907) 458; Fedtsch.; Consp. Fl. turk. Ill (l 909) 76. - Ic. : Ldb., 
Ic.pl. Fl. Ross. (1831) tab. 235; Poyarkova in Tr. Prikl. Bot., Gen. i Sel. 
XXII, 3 (1929), Fig. 42. 

Low, prostrate shrub with glabrous or spreadingly pubescent slender 
shoots; leaves 1—3 cm broad, firm, orbicular, broadly cuneate or cordate 
at base, dull or lustrous above, subglabrous on both sides and slightly 
pubescent, sometimes with scattered viscous glands, shallowly trilobate, 
coarsely dentate; petioles and inflorescence covered with spreading 
simple hairs, often with an admixture of glandular hairs; flowers dingy 
purple; berry orange -yellow, glabrous or hairy. FL May — June, 
fr. from July. 

Stony mountain slopes, among stony taluses, and in taluses in the middle 
and subalpine zones (1,500—3,000 m). W.Siberia: Alt.; Centr.Asia: 
Dzu. -Tarb., T. Sh. (E. and Centr.), Pam. -Al. (northern spurs of the Alai 
Range; in the Gissar Range and Darvaz — very rarely). Gen. distr. : 
Dzu.-Kash.: Chinese Dzungaria, Kashgariya ? Described from Bukhtarminsk 
in the Altai. Type in Leningrad. 

29. R.villosum Wall, in Roxburgh, Fl. Ind. II (1824) 514; Pojarkova in 
Acta Inst. bot. Ac. Sc. URSS, I ser., 2 (1936) 204.- R. 1 e p t o st a c hy um 
Decne. in Jacquem., Voy. d. l'Inde IV (1844) 65 (non Benth.). - R. glut i no - 
sum Decne., 1. c. (pro synon.). — R. orientale var. schugnanicum 



199 



260 



B. Fedtsch. in Trav. Mus. bot. Acad. Sc. St. Petersb. I (1902) 134.- 
R.heterotrichum A. Pojark. in Bull. Appl. Bot., Genet, a. PL Breed. XIII, 
3 (1929) 337, ex parte: quoad pi. ex Schugnan. — Ic. : Decne, 1. c, tab. 76. 

Low, prostrate shrub, profusely branching and very densely leafy; 
shoots very strong and thick, strongly pubescent, usually densely covered 
with viscous glands; petioles short, mostly half as long as blade, those on 
shortened shoots often 1 / 4 — 1 / 3 as long as blade, 1.2—3.5 cm broad, rigid, 
dull, hairy on both sides, but more densely so below, usually with a 
considerable admixture of viscous glands, trilobate or obscurely 5 -lobed, 
the lobes weakly developed, obtuse, with large, mostly subobtuse teeth; 
petioles, inflorescence axis and pedicels densely hairy; flowers purple; 
berry orange -yellow, glabrous or with sparse hairs. Fl. May — June, 
fr. July- August. (Plate XV, Figure 4). 

Stony slopes, rock crevices, taluses, from the middle mountain zone 
to 4,000 m.— Centr. Asia: Pam.-Al. — only in Shugnan, Vakhan, and 
W. Pamir (Lake Yashil-kul')- Gen. distr. : Ind.-Him., in the NW Himalayas. 
Described from Kashmir (Srinagar). Type in London. 

Note. A specimen with colorless (whitish) flowers (f. albiflorum 
A. Pojark.) was collected by I. A. Raikova on the Shakhdara River in the 
W. Pamir. 



Series 2. Alpinae A. Pojark.— Fruiting racemes usually 3 -flowered; 
flowers — including ovary — glabrous, greenish; berries red. All glands 
crystalline and stalked. Shoots unarmed, light; leaves dull, with well 
developed lobes. 

Few species distributed in the Far East, including Japan, central China, 
and Europe. 

30. R. maximoviczianum Kom., Fl. Mansh. II (1904) 787 (ex parte). - 
R.maximoviczii a umbrosum Kom., 1. c, 443. — R.alpinum J3 
manshuricum Maxim, in Mel. biol. IX (1873) 239. - R. d i s t an s a 
manshuricum Jancz. in Bull. Acad. Cracov (1906) 289; Monogr. d. 
Groseill. (1907) 460. — R.tricuspe a typic um Nakai in Bot. Mag. 
Tokyo XXX (1908) 142.- Ic. : Nakai, Fl. sylv. kor. XV (1926) tab. V; 
Poyarkova in Tr. Bot. Inst. Akad. Nauk SSSR, 1 ser., 2 (l 936 ), Fig. 15. 

Low shrub, 40—60 cm high; branches spreading, bending downward, and 
rooting, with annual brown and older brownish gray shoots; leaves on 
fruiting branchlets 1.5—4 cm long, those on sterile branches to 5.5 cm 
long, with cordate or truncate base, slender, dull, dark green and covered 
with glandular bristles above, light and glabrous or with sparse glandular 
hairs along the veins below, trilobate, sometimes with the rudiments of 
a second pair of lobes at base; lobes acuminate or acute, the middle lobe 
often rhomboid, the lateral triangular, acutely and coarsely dentate; 
staminate flowers in loose 10— 13 -flowered racemes; fruiting racemes 
1—3 -flowered; axis and peduncles beset with stalked glands; bracts longer 
than flowers, lanceolate; young berries narrowly oblong at maturity, 
oval, obovate, or clavate. Unpleasant to the taste. Fl. June, fr. August. 

Mixed forests (Siberian stone pine and broadleaf), in undergrowth and 
in the shade of rocks.— Far East: Uss. Gen. distr. : Jap.-Ch. — Manchuria 
(Kirin Province) and N. Korea. Described from the Lifuding River. 
Type in Leningrad. 



200 



(261) 




PLATE XV. 1-Ribes sachalinense Nakai.; 2-R.horridum Rupr., a) section of flower; 
3-R.saxatile Pall., a) pistillate flower, b) staminate flower; 4 - R.vill osum Wall.; 
5 -R.alpinum L.'j 6 -R.lucidum Kit.; 7 - Gros sul a ri a acicularis (Smith) Berger, 
a) flower. 



201 



263 



Note. This species should be differentiated from another Far Eastern 
representative of the subgenus Berisia — R.komarovii A. Pojark., a 
tall shrub with lustrous leaves, glabrous above, the lobes weakly developed, 
and with longer fruiting raceme; the ecological conditions also differ. 

31. R.alpinum L., Sp.pl. (1753) 200; M. B., Fl. taur. -cauc. I, 170; Ldb., 
Fl. Ross. II, 1, 196; Shmal'g., Fl. I, 357 (ex parte); Grossg., Fl. Kavk. 2 
(1930) 241. — Gros sularia insipida Rupr., Fl. ingrica (i860) 419 
(pro max. parte). - Ic: Rchb., Ic. Fl. Germ. XXIII, tab. 135. 

Shrub to 1.5 m high, with slender shoots initially brown, turning gray 
toward the second year; leaves 1.5 — 4 cm in diameter, those on sterile 
shoots slightly larger, with cordate, truncate, or rounded -cuneate base, dark 
green and covered by glandular bristles above, light and glabrous or slightly 
hairy-trilobate or with the rudiments of a scond pair of lobes at base; lobes 
below, acute, broadly triangular, the middle lobe sometimes elongated, 
rhomboid; staminate flowers 8 — 12 in racemes, 3.5 — 4 cm long; fruiting 
racemes short, 3-, sometimes 2- or 4 -flowered; pedicels of pistillate flowers 
short, 1.5 mm long, 2.5 — 3 mm in fruit, glandular like the axis; berry red, 
orbicul'"'" from the very beginning, globose at maturity, 7 — 9 mm in diameter, 
unpleasant to the taste. Fl. May— June, fr. July. (Plate XV, Figure 5). 

Broadleaf and spruce-broadleaf forests, forest edges, among shrubs, 
riverbanks; on slopes: mainly stony — limestone as well as granite slopes; 
rocky forest sites in the Caucasus.— European part: Lad. -Ilm., U. Dnp. ; 
Caucasus: Cisc, W. and S. Transc. Gen. distr.: Scand., Centr. and S. Eur. 
Described from Sweden. Type in London. 

Economic importance. Berries used locally in Europe for the preparation 
of fermented fruit drinks. Ornamental. 



Series 3. Lucidae A. Pojark.— Fruiting racemes 5 — 11 -flowered; 
flowers — including ovary — glabrous, greenish; berries red; glands 
crystalline, stalked. Shoots light, unarmed. Shrubs with lustrous leaves. 

Few distributed in the Far East (except Japan), in central China and 
Europe. 

32. R. komarovii A. Pojark. in Acta Inst. bot. Acad. Sc. URSS, ser. I, 2 
(l 936)209.— R. ma x i mo vi c z i i (m ax im o vi c z i anu m) j3 saxatile 
Kom., Fl. Mansh. II (1904) 443 (conf. p. 787).- Ic: Poyarkova, 1. c, Fig. 16. 

Branching shrub to 2.5 m high, with glabrous shoots; leaves 2 — 4.5 cm 
in diameter, firm, orbicular, with truncate or rounded- cuneate base, lustrous, 
light green, glabrous above, with sparse glandular bristles along the veins 
below, trilobate, with slightly developed obtuse lateral lobes and larger, 
sometimes acute, middle lobe; staminate flowers unknown, pistillate flowers 
in erect 5— 11 -flowered racemes, small, greenish, with oval sepals and very 
small cuneate petals, the pedicels 0.5— 2.5 mm long, covered — like the axis with 
stalked glands; berry oblong when young, globose at maturity. Fl. May, fr. July. 

Scrub, open stony slopes, rocks, crests, calcareous outcrops.— Far East: 
Uss., Suifun River basin. Gen. distr.: N. Korea (on the Yalu River). 
Described from the Vostochnyi Ushagou River, a tributary of the Suifun, 
in the [former] Ussuri Territory. Type in Leningrad. 



202 



Economic importance. Ornamental owing to beautiful lustrous foliage 
and dense rounded crown; the berries can be used to make fermented drinks. 

33. R. lucidum Kit. in Linnaea XXXII (1863) 481; Pojarkova in Acta 
Inst. Bot. Ac. Sc. URSS, ser. 1, 2 (1936) 210. - R. alpinum Jancz. in Monogr. 
d. Groseill. (1907) 460 (non L.); Shmal'g. , Fl. I, 357 (ex parte). - Ic: 
Syreishchikov, Fl. Mosk. gub. II (1907) figure on 223 (under R. alpinum); 
Poyarkova, 1. c, Figure 17. 

Shrub to 2 m high, profusely branching, with dense crown and erect 
branches; leaves 2 — 5.5 cm in diameter, to 6.5 cm on sterile shoots, oblong- 
cuneate, lustrous, usually glabrous above, less often with sparse glandular 
hairs, trilobate, the middle lobe acute, elongate, the lateral smaller and 
more obtuse, with large, sparse, mostly obtuse teeth, the blade base rounded 
or cuneate, rarely truncate at base; racemes of staminate flowers 2.5— 5.5 cm 
long, 12 — 30-flowered, fruiting racemes 5 — 8-flowered; axis and pedicels 
glandular, the latter short in pistillate flowers, 0.5 — 1 mm in fruit 2.5 — 3 mm 
long; berry initially oblong, subglobose at maturity, 7— 10 mm in diameter, 
red, unpleasant to the taste. Fl. May — June, fr. July. (Plate XV, Figure 6). 

Slopes in shrub thickets and forests.— European part: Lad. -Ilm., U. 
Dnp.; Caucasus: W. and E. Transc. Gen. distr.: Scand., Centr. and S. Eur. 
Described from Hungary. Type unknown. 

Note. R. lucidum Kit. occurs apparently more frequently in the 
European part of the Soviet Union than R. alpinum L., but is rare in the 
Caucasus. There are herbarium specimens from two sites, from the 
vicinity of Bakuriani and from Circassia on the Black Sea coast. In the 
European part, it forms hybrids with R. alpinum L. which either possess 
characters intermediate between the 2 species or else are closer to 
one of them. 

Economic importance. As for R. alpinum L. 



Series 4. Diacanthae A.Pojark.— Flowers greenish, glabrous, including 
ovary; leaf lobes weakly developed; shoots light brown, usually with paired 
prickles in the nodes and often with spiny internodes. Pubescence short, 
velutinous; all glands stalked. 2 species. 

34. R. diacantha Pall., Reise III (1776) 722; Ej., Fl. Ross. I, 2 (1788) 36; 
Turcz., Fl. baic. -dah. 1, 441; Ldb., Fl. Ross. II, 1, 196; Kom., Fl. Mansh. II 
(1904) 445; Kom. and Alis., Opred. rast. Dal'nevost. Kraya I (1931) 621.- 
Ic. : Pall., Reise III, t. I, fig. 2; Fl. Ross, tab. 66. - Exs.: Herb. Fl. URSS No. 
3140. Vernacular name: taranushka. 

i Shrubs to 1 m high; branches very numerous, fascicular, sometimes 
nodding; leaves 1.5 — 3 cm long, thin but firm, glabrous on both sides, dark 
green and lustrous above, light and whitish below, with clearly delineated 
but not prominent veins, oblong- cuneate, shallowly trilobate or with lobelike 
incisions in upper part, the lobes with few acute apiculate teeth on the 
margin; inflorescence glabrous; berry red, small, 5— 7 mm in diameter, 
globose, not edible. Fl. May— June, fr. July. Stony slopes, rocks, taluses; 
usually in small thickets. — E. Siberia: Dau. Gen. distr.: Mong. (NE Mong.) 
and Jap. -Ch. (NW Manchuria (near Chailar) and N.Korea). The Korean 
specimen is distinguished by the presence of very large, long- stalked glands 
on leaf margins. Described from Dauria. Type in London. 



203 



266 



Economic importance. Ornamental plants. 

35. R. saxatile Pall, in Nov. Acta Acad. Petrop. X (1797) 376; Ldb., PI. 
Ross. II, 1, 195; Pojarkova in Bull. Appl. Bot., Gen. a. Plant Breed. XXII, 3 
(1929)336; Kryl., Fl. Zap. Sib. ; VI (1931) 1447. - R. c une at urn Kar. et 
Kir. in Bull. Soc. Nat. Mosc. XIV (1841) 426; XV (1842) 356. - R. d ia c a nt h a 
var.typica Trautv. in Bull. Soc. Nat. Mosc. XXXIX, 1 (1866) 314. -R. dia- 
cant'ha (non Pall.) Rgl. et Herd, in Bull. Soc. Nat. Mosc. XXXIX, 2 (1866) 
68; Kryl., Fl. Alt. II (1902) 463.- Ic: Ldb., Lc.pl. Fl. Ross. (1831) tab. 239; 
Poyarkova, 1. c, Fig. 41. Vernacular name: taranushka. 

Low, spreading shrub, 60 — 90 cm high; leaves 1—2.5 cm long, firm, 
glaucous -green, glabrous on both sides when adult, less often puberulent 
below, rounded-obovate, with broadly cuneate base, shallowly trilobate at 
the apex, with 3 large lobelike teeth, the lobes obtuse or subacute, with 
1 or 2 acute teeth on the margin; inflorescence axis and pedicels densely 
puberulent; berry initially red, dark cherry-red at maturity, 5 — 6 mm in 
diameter. Fl. June, fr. August. (Plate XV, Figure 3). 

Rocks, taluses, among stony taluses, open slopes, also on open rocky river 
banks. — W. Siberia: Irt.,Alt.; Centr.Asia: Ar. -Casp. (western border 
passes along the Mugodzhar hills), N. Balkh., Dzu. -Tarb., T. Sh. (Mt. Ketmen). 
Gen. distr.; Dzu. -Kash. - NW China (Kuldja). Described from Dzungaria. 
Type in London. 



Series 5. Pulchellae A. Poj ark.— Flowers greenish, the sepals red on the 
margin; leaf lobes glabrous, well developed. Shoots light brown, with 
paired prickles at base of leaves and sometimes with spiny internodes. 
Pubescence crisp; all glands stalked; 1 species. 

36. R. pulchellum Turcz. in Bull. Soc. Nat. Mosc. V (1838) 191; Fl. baic. - 
dah. I (1842) 442; Ldb., Fl. Ross. II, 1, 196; Maxim, in Mel. biol. IX (1873) 24; 
Kom., Fl. Mansh. II (1903) 446; Kom. and Alis., Opred. rast. Dal'nevost. 
kraya I (193.1) 621.- Ic: Trautv., PL imag. et descr. fl. Ross. (1844) tab. 8. 

Shrub 1 — 2 m high; leaves 0.8 — 3.2 cm long, dull, glabrous or with glandular 
bristles above, trilobate; lobes acute, with simple and incised apiculate teeth; 
base truncate or rounded-cordate; inflorescence axis and pedicels glabrous 
with a pubescence of crispy hairs and an admixture of stalked glands; berry 
red, 5 — 6 mm in diameter. Fl. June, fr. August. Stony and pebbly slopes 
in the forest and steppe zones.— E. Siberia: Dau. (Selenga River basin). 
Gen. distr.: NE Mong., Jap. -Ch. N. Manchuria (Great Khingan) and N. China 
(Chihli Province). Described from Mongolia. Type in Leningrad. 

Economic importance. Graceful ornamental shrub. 

Among the species from foreign countries, the following are the most 
widely cultivated in the Soviet Union: 

*R. sanguineum Pursh, Fl. Amer. sept. (1814) 164. — Coreosma 
sanguine a Spach, Hist. d. veg. phaner. VI (1838) 155 (subgenus 
Calobotrya Spach). 

Erect shrub 1 — 3 m high, unarmed, with reddish, pubescent, sometimes 
glandular shoots; leaves firm, dark green above, pubescent, whitish- 



5773 204 



tomentose below, cordate or reniform, 3-, rarely 5-lobed, with broad obtuse 
lobes; flowers in erect, long, 10 — 20 -flowered racemes, sometimes with 
large red bracts exceeding the pedicels; bracts, pedicels, and inflorescence 
axis pubescent and covered with glands; flowers bright pink or red, large 
(to 15 mm long), with pubescent, campanulate, cylindrical hypanthium and 
oblong ovary densely covered with long- stalked glands; berry black, 
glandular. 

Native to the mountains of western North America from British Columbia 
to N. California. 

Economic importance. Ornamental. Known in a number of garden forms, 
with white, dark red, and double flowers. 

*R. aureum Pursh, Fl. Amer. sept. (1814) 164. — R. f la vu m Berl.; in 
Mem. Soc. Gen. Ill, 2 (182 6) 60. — R. t e nuif lo r u m Lindl. in Trans, hort. 
Soc. Lond. VII (182 8) 242. — Chry sobot ry a Lindleyana Spach, Ann. 
Sc. Nat. II ser., IV (1835) 20 (subgenus Symplocalyx Berl.). 

Shrub ca. 2 m high, with red young shoots; leaves to 5 cm long, 6 cm broad, 
glabrous, rounded-reniform or obovate in outline, cuneate, rounded, or 
slightly cuneate at base, trilobate, the lobes subobtuse with few teeth on the 
margin; racemes erect, 3— 7 cm long, 5 — 15 -flowered, with large bracts 
exceeding pedicels; flowers yellow, aromatic; hypanthium tubular, slender, 
5 — 9 mm long, ca. 1 .5 mm in diameter; sepals spreading at anthesis, 5 — 8 mm 
long, erect in fruit; petals half as long as sepals, orange -red. Fruits 
globose, brown-red or black, sometimes yellow (var. chrysococcum 
Rydb.). 

Native to the mountains of western North America. 

Economic importance. At anthesis a very ornamental shrub bearing 
tasty fruit. 

Avery closely related cultivated species is R.odoratum Wendl. 
[P. long if lor um Nutt., R. f r a g r a n s Lodd.,R. aureum auct. pi. (non 
Pursh)], distinguished from R. aureum Pursh by the following characters: 
larger flowers, hypanthium 12 — 15 mm long, sepals 5 — 6 mm long at 
anthesis, recurved or spreading (not erect! ) in fruit, larger and usually 
deeper-lobed leaves. Berries black or orange-yellow.— North America, 
in the plains from the Rocky Mountains to Minnesota and Arkansas, as far 
south as Dakota and Texas. 



Genus 714. GROSSULARIA MILL. 

MilL.Gard. Diet. ed. 7 (1759). 

Shrubs with 3- or 5-, less often 7 -partite or solitary spines at nodes of 
shoots and mostly with spiny internodes, with alternate, palmatilobate, 
dentate leaves; racemes fascicular, usually 1 — 3 -flowered; pedicels not 
developing; flowers bisexual, 5-merous, with campanulate hypanthium, 
oblong recurved sepals, and small petals; style 1, split usually not beyond the 
middle; ovary pedicels long, usually many times as long as ovary, not 
disarticulating from fruit; fruit a succulent berry with dried-out flower, 



* From the French grosei 1 1 e, currant. 



205 






,u tin- apex Like the ovar) smooth, hairy. glandular hispid or spiny; 

Seeds With hard endoplourn .iu*l gelatinous testa. 

i. Spines .11 the nodes 7; ovary and berry spiny 

3. C.. burejensis (l<Y. Sehmidl ) Iht ; mt. 

* Spines .u the nodes tripartite; berry smooth or glandular hispid, 

luit noi spiny 2. 

i eaves and Flowers glabrous; berrj always smooth: spinos at tin- 

Lnternodes persisting 3 years or more 

2. G.acicularis (Smith) Spach. 

■+ Leaves pubescent or glandular hispid; flowers pubescent; berrj 

glandular or smooth; only Lnternodes of annual shoots spiny 

' 1. G. reclinata (I .) Mill. 

i. G. reclinata (L.) Mill, in c,,u\\. met. ed. 8 (1768). Ribes 
reclinatum L., Sp.pl. (1753) 2Q1, R.grossularia L., 1. c; Ulb., 
i'i. Ross. ii. i . L94; Shmal'g., PI. l. 35; Grossg., PL Kavk, u (1930) 241.- 
R.spinosum Gilib., PL Lithuan, u U 785) L 3-t . u. u v a c r is pa rail.. 

FLROSS. I, 2 (1788)37 (uon 1,.). K.caucasicum Adams in Roem. et 
Schult., Syst.veg, "\ (1810) 507. u rossu la r ia vulgaris Spech, 
Hist. veg. ph. in. V] (1838.) 171. K.vulgare *.\ Koch in Linnaea XVI 

(1842)355 (non Lam.). Gr.spinosa Rupr., i'i. ingricn (1860)415.— 
[c: Berger Ln N. York St. Agric, Exp, St, rechn. Bull., No. L09 (192 I ) 

pi, \ in. 

Shrub ca, i m bigh; spinos at the nodes to 1 1.4 cm Long, those on 
Lnternodes short, mostly rather sparse, sometimes absent: leaves 
1 — 3 . t> (6)cm bro.nl. 3 or 5 -lobed, dull, puberulenl on both sides, obtusely 
dentate: flowers solitary or m pairs, pubeseent. greenish, sometimes 
reddish; ovary more or Less glandular hispid (var.glanduloso - 
setosa W.Koeh ), producing glandular-hispid or nearly smooth berries, 
or else smooth (var. glabra YV. Koeh): berry globose or broadly ellipsoid. 
greenish or yellowish, usually more or loss reddish to dark red. PL May, 
,lune. t'r. July. 

Among shrubs on slope in the middle mountain zone.— European part: 
wild only in M. Dnp. (w. Ukraine), occasionally wild in : Lad.-iim.. v. v., 
U.Dnp., V.-Don; Caucasus: wild in: Cisc, Hag., \V., K.. and S. lYansc. 
Gen. distr. : Centr.and S, Eur., including Great Britain and N.Spain; 
mountains of N, Afr. Described from Germany. Type in London. 

Note. Linnaeus described 3 species of Kuropcan currants considered 
by recent authors as es of a single species: R. reclinatum L. 

(R, g r o s s u I a r i a v. r cell n a t u m Mori., K. g. y g I a b r u m \V. Koch). 
R, g r S s u la r i a l. U\- g. ' ! g ' ^ n d u I o s o -set o s u m W. Koch) and R.uv a 
c r 1 s p a I . (<d r o ssul a r i a u v a c r i s p a Mill.. K. g. p p u b c s c e n s 
W. Koch.. R. g. v. uv a oris pa Jancz.). 

G.uva crispa td .) Mill. There arc grounds for considering this an 
., .. independent species native only to S. and \\ . Europe. It is distinguished 
by ovaries covered With a dense pubescence of simple hairs, small, 
yellowish, pubescent berries, smaller, densely pubescent leaves, absence 
ofacicularspuics.it Lnternodes of shoots, smaller size, and later (by - weeks) 
flowering. In the Soviet Union. G.uva crispa occurs only in cultivation 



:oo 



in a number of varieties yielding small, very sweet, pubescent berries, 
or as an escape. As regards the 2 other Linnaean species, lack of 
material has prevented elucidation of their interrelation; it may be 
coi rect to group them, as Spach did, in one species. The fact, is that the 
distinguishing character — the glabrous or glandular -hispid ovary — is n 
constant, showing a series of transitions. The distribution of these forms 
within the- limits of the range is not uniform: th< ilar -hispid form is 

frequent in Europe, whereas specimens with glabrous oval r rarely; 

in the Caucasus, however, there are almost exclusively specimens with 
glabrous ovary. Clarification is also needed as to whether the Caucasian 
( ur rant is identical with the European in other characters. 

Economic importance. This is the ancestor of most cultivated varieties 
of currants. The berries are eaten raw or are used for making jam, 
syrups, jelly, and fruit, wine. Chemical composition of the berries, according 
to Wehmer (percent): water £5.4%, invert sugars 4.67, saccharose 0.41, 
free acids 1.9 (malic), tannic acid 0.089, nitrates 0.94, raw cellulose 
2.7, ash 0.51; the ash is rich in alkali (ca. 49%) and phosphoric acid (10,68%); 
the seeds contain ca. 1 9.8% of fatty oils; according to Hegi, the berries 
contain (percent): 84 -88% water, 8- -8 dextrose and levulose (not containing 
saccharose), 0.3— 0.57 nitrogen, 1 .13 pectins, 3.52%, cellulose, 1 .42% free- 
acids (citric, tartaric, and malic; unripe berries also contain succinic 
acid), ash, and a small quantity of tannic acid and resin. 

2. G.acicularis (Smith) Spach, Hist. veg. phan. VI (1838) 173; Berger 
in N.York. Agric. Exper. St. Techn. Bull., No. 109 (1924) 107; Poyarkova, 

in Tr.Bot. Inst. Akad. Nauk SSSR, 1, ser. 2 (1936) 214.- Ribes acicularis 
Smith in Rees, Cyclop. XXX (1819) 372; Ldb., Fl. Ross. II, 1 ; 194; Kryl., 
FL Zap. Sib. VI (1831)1446.- Ic: Ldb.; Ic, Fl. Ross. (1831 ) tab. 230. 

Shrub; shoots densely covered with slender, acicular spines; spines 
at the nodes to 1 cm long; leaves 0.7—3 (3.5) cm broad, glabrous, lustrous 
below, 3- or 5-lobed, with acute duplicato -dentate lobes; flowers solitary, 
greenish white or pinkish, glabrous; ovary and berries smooth, the latter 
12—15 mm in diameter. Fl. June, fr. July. (Plate XV, Figure 7 J. 

Open stony mountain and hill slopes.— W.Siberia: Alt.; E.Siberia: 
W. part of Ang.-Say.; Centr. Asia: Dzu.-Tarb.; the SW border passes 
through the Dzungarian Ala-Tau. Gen.distr.: Mong. (NW and Centr.). 
Described from the Altai. Type in London. 

Economic importance. Distinguished by its large tasty berries; 
deserves the attention of fruit growers. Berries are edible raw or 
can be used for the making of syrups, jams, and wine. 

3. G.burejensis (Fr. Schmidt) Berger in N.York Agric. Exper. St. 
Techn. Bull., No. 109 (1924) 112.- Tibes burejense Fr. Schmidt in 
Mem. Acad. Sc. St. Petersb. VII, ser. XII, 2 (1863)42; Maxim, in Mel. biol. IX 
(1873)216; Kom., Fl. Mansh. II (1903) 435; Kom. and Alis., Opred. rast. 
Dal'nevost.kraya I (1931) 621.- Ic: Fr. Schmidt, I.e., tab. I, f. 1. 

Shrub; shoots very spiny, densely covered with large and small spines; 
spines at the nodes to 1 cm long; leaves 1.5— 5 cm long, to 8 cm broad on 
sterile shoots, deeply 3 - or 5 -lobed, puberulent on both sides or with 
glandular bristles only along the veins below, the margin acutely incised - 



207 



dentate; flowers large, pinkish, always solitary; ovary hispid; berry 
spiny, dingy green. Fl. May, fr. July. 

Mountain coniferous forests, slopes, stream banks; solitary or in small 
groups.— Far East: S. part of Ze.-Bu.,Uss. Gen. distr. : Manchuria and 
N. Korea. Described from the Bureya River. Type in Leningrad. 



Family * PITTOSPORACEAE LINDL.* 

Flowers bisexual, more rarely diclinous, solitary or in a corymb. 
Sepals 5, free or fused below, imbricate; petals 5, free or slightly fused 
at the base. Ovary superior, 2—5 -locular. Style simple, with capitate 
stigma. Fruit usually a capsule, less often a berry. Embryo small, 
with copious endosperm. Trees, shrubs, less often climbing, sometimes 
spiny plants with erect, glabrous, spiral or alternate leaves. 

Fossilized P it t o s po ru m (Sapotacites)Putterlicki Ung. 
occurs in lower Pliocene deposits of W. Transc, in volcanic tuffs of the 
Goderdzi Pass. 

Genus * PITTOSPORUM ** DRYAND 

Dryand ex. Gaertn. Fruct.I (1788) 296, tab. 59. 

Sepals free or fused at the base. Petals approximate at the base or to 
the middle or else fused, usually recurved at the apex. Stamens subulate 
with ovate-oblong anthers. Ovary sessile or subsessile, 2 -, less often 
3— 5 -locular, with capitate stigma. Capsule globose, ovoid, or oblong, 
2 - or 3 -valved, with parietal placentation and coriaceous, almost woody 
p 71 lobes. Seeds glabrous, agglutinated by a resinous substance. Shrubs, less 
often small trees, the more or less evergreen leaves in some species at 
tips of shoots. Flowers in terminal or lateral inflorescences, in a corymb, 
umbel, or panicle, less often solitary. 

*R.tobira t Dryand in Ait. Hort, Kew. 2 (1810)27.— Ic: Bot. Mag. tab. 
1396 (1810); Nicholson, Diet, of Gardening III (1867) 154, f. 193.- Pax in 
Engl.u. Pr., Nat. Pflzf. Ill, 2a (1891 ) 110, f. 72. 

Low shrub, profusely branching; leaves spirally arranged, coriaceous, 
obcuneate, rounded at the apex, entire dark green, mainly at apices of 
shoots; flowers in dense terminal inflorescences; calyx pubescent, with 
ovate sepals; petals obcuneate, recurved, rounded at the apex, cream- 
colored or pale yellowish, aromatic; short filaments; ovary bottle -shaped; 
style short; capsule acuminate, many -seeded; seeds reniform, slightly 
acuminate unilaterally. 

Often cultivated in gardens in Transcaucasia and on the south coast 
of the Crimea; satisfactorily naturalized in the Mediterranean area and 
S. Europe. Native to the coasts of Japan. 

Economic importance. Cultivated as an ornamental shrub; flowers 
aromatic, can be used for perfumery. 

* Treatment by I. V.Palibin. 
** From the Greek pi tta , pitch and sp or a, seed. 
t From tobera.the Japanese name of the plant. 



208 



Family LXXV. HAMAMELIDACEAE LINDL.* 

Flowers bisexual, less often unisexual, actinomorphic or zygomorphic; 
corolla sometimes irregular, adnate to ovary. Petals 5 or more; stamens 
6—7, free. Ovary bilocular, superior; locules with 1 to several ovules. 
Styles subulate. Fruit a bilocular capsule. Seeds with fleshy endosperm 
and erect embryo. Trees or shrubs with sparse, stellate pubescence. 
Leaves usually deciduous; stipules paired. 

The following representatives of this family are known from fossile: 

Parrotia fagifolia Goepp., in Lower Pliocene of the Goderdzi Pass 
(W. Transc. ); P.pristina Ett. in L. Don, Lower Sarmatian (Krynka). 

Liquidambar europaeum A. Br. in Upper Oligocene of 
N. Kazakhstan: Dzharkue, Bol'shie Barsuki, Kara-Sandyk, Sarry-Bulak, 
Chushka-kul' and Mynsai (indricotherium horizon). Upper Oligocene 
deposits in the vicinity of Tomsk. Sterlitamak district, Lower Miocene, 
Shkatla River. Sakhalin Island, Upper Due deposits (Endo). Vicinity 
of Vladivostok: Upper Due deposits. 

Fothergilla ungeri Kov. — in Sarmatian of Bl. (Amvrosievka). 



Genus 715. PARROTIA ** C. A. M. 

C.A.M.Verz.(1831) 46. 

r 

Flowers bisexual, capitate; bracts broad, membranous, resembling 
an involucre. Calyx campanulate, with 5—7 -lobed limb, coriaceous; lobes 
foliiform. Petals absent, stamens 6—7, opposite to sepal lobes; filaments 
long; anthers oblong, tetragonal; ovary half -inferior, the locules usually 
1 -seeded; ovules pendulous; styles filiform, simple; fruit dehiscing 
by 2 valves. 

1. P.persica (DC.) C. A. M., 1. c; Boiss., Fl. Or. II (1872) 818; Medv., 
Der. i Kust. Kavkaza, Grossg., Fl. Kavk. II (1930) 242. — Hamamelis 
per sic a DC, Prodr. IV (1830) 268. - Ic. : Bot. Mag. (1868) 5744; 
Volf and Palib., Opred. (l 904) 336-337.- Exs.: Hohenacker, Lenkoran, 
No. 2117; PL orient exsic, No. 388; Herb. fl. cauc, No. 224. Vernacular 
name: zheleznoe derevo. 

Tree; branches angular, often connate when adult, covered with stellate 
pubescence when young; leaves short -petioled, cuneate at base or rounded - 
ovate or obovate, slightly oblique, entire in lower part, emarginately and 
coarsely dentate in upper part, glabrous above, slightly pubescent with 
sparse, stellate hairs; flowers appearing before leaves in 2 - or 3 -flowered 
fascicles or heads, short -pediceled; bracts membranous, large, ovate -oval, 
covered on the outside with dark rufous -brown tomentum, glabrous inside; 
calyx broadly campanulate, with 6 or 7 remote, obtuse, foliiform sepals, 
ciliate at the apex; tube covered with stellate hairs; filaments long; 
anthers acuminate, brightly colored; capsule woody, rounded -oval, the 



Treatment by I. V. Palibin. 

Named after I. F. Parrot, a Russian naturalist (1791- 1841) who climbed Mt. Ararat and ascertained 

its altitude. 



209 



273 



valves elongate -acuminate, reflexed, much longer than calyx; seed ovoid, 
acute, with 2 scars at base, pale yellow. Fl. March, fr. September. 
(Plate XVI, Figure 3). 

Caucasus: Talysh forests. Gen. distr.: N. Iran (Gilan and Mazanderan). 

Economic importance. Very sturdy wood with many applications; 
produces high-quality coal; branches used for enclosures and fences. 
Certain representatives of this family are beautiful shrubs flowering in 
early spring before the leaves bloom; cultivated in humid subtropical 
regions (Sukhumi-Batumi). The following are related: Corylopsis 
pauciflora S. etZ., C.spicata S.etZ. (both from Japan), Sycopsis 
sinensis Oliv. (China); Hamamelis japonica S. et Z. (japan), and 
H. virginiana L. (N. Am.) — the leaves of the latter are used in medicine; 
Liquidambar styraciflua L. (N. Am.), cultivated in gardens on the 
Black Sea coast of the Caucasus and the Crimea. 



Family * EUC0MM1ACEAE VAN-TIEGHEM* 

Flowers unisexual, without perianth; staminate flowers in fascicles, 
numerous (4— 10 ), narrowly linear, to 1 cm long; pistillate flowers solitary, 
short -petioled; ovary flat, bilocular, ovules 2; fruit a nondehiscent, 
unilocular capsule, elongate -ovoid; seed 1, lanceolate. 

Occurs in Sarmatian deposits of the Bl. (Krynka, Orekhov). 



Genus *E0C0MMIA* OLIV. 

Oliv. in Hook., Icon. pi. (1890) tab. 1950. 

1. E.ulmoides Oliv., 1. c; Harms in Engl. u. Pr., Nat. Pflzfam. 1 Aufl. 
Nachtr. I (1897) 159; Nachtr. II (l 908) III; 2 Aufl. Bd. 18a (l 930) 351; 
C. I. Schn., Ill, Handb. Laubhk. I (1904) 424; Wilson in Sargent, PI. Wilson. 
1.3 (1913) 433.- Ic: Hook., 1. c, t. 1950, tab. 2361 ; C. I. Schn., 1. c, f. 2 70; 
Silva Tarouga et C. K. Schn., Freil. Laubh. (1922) 198, Abb. 223; Hutchinson, 
Fam. Flowering pi. (1926) 215, f. 182; Harms (1930) 351, f. 182.- Exs.: 
A. H. Steward, PL Kweichow (l 930), No. 41 7. 

Tree 20—30 m high, with dull dark gray bark; leaves herbaceous, 
elongate -elliptic, obtusely narrowed toward base and elongated toward 
the apex, glabrous above, slightly pubescent along the veins below, denticulate, 
the leaf teeth recurved near the apex; petioles glabrous, slender, % — x / 5 
as long as the blade; fruits unilocular, in short fascicles, winged, elongate - 
elliptic, coriaceous, lustrous, with a notch at the apex; seeds narrowly 
lanceolate, with hard testa. 

Grows in mountainous regions of W. and central China. In the Hupeh, 
Szechwan, Shansi, and Kansu provinces; cultivated in European parts, 
where it withstands frosts of 15— 20°C; cultivated in the Soviet Union in 
the subtropical regions of Transcaucasia (mainly in Abkhazia) as an 



Treatment by I. V. Palibin. 

From eu.real and ko m ma.hair, referring to the hairlike appearance of the gum filaments visible when 

the bark and leaves are torn. 



210 



economically important tree yielding gutta-percha (see V.N. Andreev 
"Evkommia, kitaiskoe guttaperchevoe derevo na Ukraine i na Kavkaze" 
[Euc omm ia, the Chinese Gutta-Percha Tree in the Ukraine and the 
Caucasus]. Gos. Trest Kauchukonos. Ukrainskii Nauchno-issledovatel'skii 
Institut Kauchuka i Kauchukonosov ["Kauchukonos" State Trust. Ukrainian 
Scientific -Research Institute or Rubber and Rubber-bearing Plants]. 
Kiev 1932, p. 80). 

Economic importance. This tree has been known in China as medicinal 
since ancient times under the name "Dzu-Dzhun." The bark and leaves 
contain a gutta-percha substance, in the form of silvery filaments — 3% of 
dry weight. The tree is grown in the subtropical regions of Transcaucasia 
for the extraction of gutta-percha. 



Family LXXVI. PLATANACEAE LINDL.* 

Flowers unisexual, in globose inflorescences, with long pendent peduncles 
developing at tips of current -year shoots. Calyx absent. Staminate 
inflorescences: compact, rounded or elongated, with 3—8 staminate flowers; 
stamens subsessile, the anthers bilocular, corymbiformly truncate at the 
apex, mixed with fleshy cuneate scales. Pistillate inflorescences: numerous 
sessile flowers surrounded by 3—8 cuneate obconical scales; ovary 
unilocular, with long subulate apically curved pistils; ovules pendulous, 
erect. Fruit a nondehiscent nut surrounded at base by slender articulate 
hairs, 1 -seeded, with erect, linear embryo. Trees with green-gray bark, 
separating in large flakes, the alternate palmatilobate or lobate 
leaves covered when young with stellate hairs; stipules deciduous. 

Representatives of this family — Credneria, Protophyllum (?), 
etc.— and even of the genus Platanus appear as early as the Middle and 
Upper Cretaceous, in Ar.-Casp. and V. -Kama, the Caucasus, and the 
Far East, Sakh., Kamch., Okh., An., Ang.-Say., Ze. -Bu. — Platanus 
aceroides Goeppert in the Tertiary (Sarmatian) of Bl. (Krynka). — 
P.guillelmae Goeppert in the Tertiary of Sakh. (Pil'vo), Uss. (De Friz 
Peninsula), Balkh. (Ashutas). 



Genus 716. PLATANUS** L. 

L., Sp.pl. ed.l (1753) 998. 

The generic characters of the genus appear in the description of the 
(single genus) family. 

1. Leaves slightly lobate, angular, with inconspicuous teeth; heads 1—3 

on inflorescence axis; fruits clavate, glabrous, with minute style .... 
3. P. occidentalis L. 



Treatment by I.V.Palibin. 

From platanos, the name of this tree in ancient Greece. 



211 



275 



276 



+ Leaves deeply lobed, more or less dissected, with conspicuous teeth; 

heads elongated, usually pubescent 2. 

2. Heads up to 6, small (to 1—1.5 cm in diameter); leaf lobes with 
numerous large and small teeth; fruit glabrous at the apex, with 

long straight style *P. orientalis L. 

+ Heads large, to 2—2.5 cm in diameter 3. 

3. Leaves on branches 3— 5 -lobed, with cordate base; fruits subobtuse, 
covered with whitish lanate hairs; style short *P. cuneata Willd. 

+ Leaves palmatilobate, slightly dentate, pubescent with long hairs; 

style long 4. 

4. Leaves on branches truncate at base; fruits acuminate, pubescent 

with rigid hairs; style somewhat curved, 3 / 4 as long as fruit 

1 . P. digitata Gord. 

+ Leaves on branches with very remote deep few -toothed lobes; fruits 
subobtuse, pubescent; style as long as fruit, curved, often uncinate . . . 
2. P. orientalior Dode. 



Section 1. ORIENTALES Dode, Note dendrol. (1908) 51.- Leaves deeply 
dissected, dentate; heads numerous. Fruits elongated, acuminate, pubescent, 
produced into a long style. 

*P. orientalis L., Sp. pi. ed. I (1753) 998; Ldb., Fl. Ross. Ill (1846-1851) 
595 ex pte; Boiss., Fl. Or. IV (1879) 1161 ex pte; Dode, Note dendrol. (1908) 
92; C. K. Schn., Handb. Laubholzk. II (1912) 957; Hegi, III, Fl. Mitteleuropa 
IV, 2 (1925) 661.- P. or ientalis var. insular is DC, Prodr.XVI, 2, 
(1864) 159 (excl. pi. cret.). - Ic. : Dode, 1. c; Elwes and Henry, Tr. Gr. Brit, 
a. Irel. Ill, 1, 15, tab. 204, f . 4 (1908); Henry a. Flood, Proceed. R. Irish Acad. 
XXXV B, No. 2, tab. V, f . 1 ; Hegi, 111. Fl. Mitteleur. IV, 2 (1925) f. 994.- Exs.: 
Bourgeau, Pl.de Rhodes (1870), No. 146; Peyron, Fl. Syr. (1881 ), No. 1869. 

Tree; leaves 5 -lobed, deeply divergent on shoots, with numerous acute 
teeth and with cordate base; leaves on trunks 3— 5 -lobed, more or less 
truncate, lobate or dentate at base; leaves on branches deeply divergent, 
slightly lobate and slightly dentate, with broadly cordate base; heads 
3—5, sometimes 6, small (to 1.5 cm in diameter), lateral heads sessile or 
pedunculate; fruits thickened toward the apex, pubescent, produced into a 
long, erect style. 

Not growing wild in the USSR, sometimes cultivated in gardens of 
Transcaucasia (Tbilisi). Native to southern Asia Minor, the island of 
Rhodes, and Syria; also cultivated in Palestine and other areas in the 
Mediterranean region. 

*P. cuneata Willd., Sp. pi. IV (1805) 473; Dode, Note dendrol. (1908) 54; 
C. K. Schn., Handb. Laubholzk. II (1912) 957; Hegi, 111. Fl. v. Mitteleuropa IV, 
2 (1925) 658.- Ic: Dode, 1. c, 55; Henry a. Flood, Proceed. R. Irish Ac. 
XXXV B, No. 2, tab. VII, f. (1919) 6.- Exs.: Fl. ital., No. 780 (Sicilia); 
Bornm., PL Lydiae et Cariae, exs., No. 9932; Sint. et Bornm., It.turcz., 
No. 1467. 

Tree; leaves on shoots 5 -lobed, not very widely divergent, the lobes 
and teeth numerous, with broadly cordate, subtruncate base; leaves on 
trunk 3— 5 -lobed, often trilobate, cuneate, with strongly dentate lobes at 



212 




PLATE XVI. 1 — Platanus d i git a ta Gord., leaf and inflorescence, a) mature fruit; 2 — P. orient ali or 
Dode, leaf and inflorescence, a) mature fruit; 3— Parrotia persica C.A.M., a) flowering branch, 
b) stamens, c) ripe capsule, d) young ovary, e) flower, f) fruit (Figure 3, according to Hooker). 



213 



base; leaves on branches 5-lobed, slightly dentate, truncate, broadly 
rounded at base; heads up to 6 but usually 3— 5, large (to 2.5 cm in diameter), 
the lateral sessile or petiolate; fruits thickened, somewhat obtuse, densely 
pubescent with slender whitish hairs; style short. 

Not growing wild in the USSR; cultivated in parks and gardens in the 
northern part of the Black Sea coast of the Caucasus and of the Crimea, 
sometimes also of Central Asia (Turkm.). Native to Asia Minor (N.part), 
Macedonia, Greece, S. Italy, and Sicily. 

1. P. digitata Gordon, the Garden 1 (1872)572-573; Dode, Note dendrol. 
(1908) 58; C. K. Schn., Handb. Laubholzk. II (1912) 957; Hegi, 111. Fl. v. 
Mitteleur. IV, 2 (l 92 5) 658. - P. o r i e nt a 1 i s Ldb., Fl. Ross. Ill (1846-1851) 
595, Non L. - Ic: Sibth., Fl. graeca (l 840) tab. 945 ; Dode, 1. c, 59, 

Henry a. Flood, Proceed. R. Irish Acad. XXXV, B, No. 2, tab. 8, f . 7 (1919). — 
Exs.: Fl. graeca, exs., No. 1174; Sintenis, It. troj. (l 883), No. 815; Aucher 
Eloy Herb. d'Or., No. 5330. 

Tree; leaves on shoots with 5 deeply lobed, slightly cordate; leaves 
on trunk 3— 5-lobed, cuneate at base, more or less lobate and dentate; 
leaves on branches 5-lobed, slightly dentate, truncate at base; heads 
large (to 2 cm in diameter), up to 5, but usually 2—4 and then smaller, 
sessile; fruits thickened, tapering to long, rigid, suberect style, sometimes 
recurved in a loop. April. (Plate XVI* Figure 1 ). 

Caucasus: E. and S. Transc; often cultivated in parks and gardens. 

2. P. orientalior Dode, Note dendrol. (1908) 57-58; C. K. Schn., Handb. 
Laubholzk. II (1912) 957; Hegi, 111. Fl. Mitteleur. IV, 2 (1925)658.- 

P. or i en talis Brandis, Ind. Trees (1907) 619, non L. — Ic. : P. orientalis 
Pall., Fl. Ross. II (1788) tab. 51; Dode, 1. c. - Exs.: Sintenis, Iter Transc. - 
pers.(l900) No. 1062; Herb. Schlagintweit, Kashmir, No. 14179; Bornm., 
It. pers.-turc, No. 451 8; Aitchison, Afghanistan, No. 259. 
-, 7q Tree; leaves on shoots deeply 5-lobed with numerous teeth and cordate 

lobes, base cordate; leaves on trunk 5-lobed, cuneate, markedly lobate and 
dentate at base; leaves on branches 5-lobed, subentire, rarely dentate 
and deeply parted, cordate at base; heads 3—4, less often 6, large (to 2.5 cm 
in diameter) sessile or pedunculate; fruits acuminate, the slender style often 
looped -recurved, as long as fruit. March — April. (Plate XVI, Figure 2). 

Centr. Asia: Mtn. Turkm., Amu D., Syr D., Pam.-Al. Gen. distr.: As. Min. 
N. part), N. Iran, Afghanistan, N. India (Kashmir). Undoubtedly growing wild 
in southern Central Asia, occurring in parks and gardens in more northerly 
districts. 



Section 2. OCCIDENTALIS Dode, Note dendrol. (1908) 51. - Leaves 
lobate, slightly dissected, acutely denticulate; heads 1—3. Fruits clavate, 
glabrous, with minute style. 

*3. P. occidentalis L., Sp. pi. ed. I (1753) 999; Sargent/Sylva of N. America 
VII (1895) 326-327; Dode, Note dendrol. (l 908) 65; C. K. Schn., Handb. 
Laubholzk. II (1912) 957.- Ic: Sargent, 1. c, tab. 326-327; Dode, I.e., 66; 
Henry and Flood., Proceed. R. Irish Acad. XXXV, B, No. 2 (1919) tab. V, f. 2, 
tab. IX.- Exs.: Somes, Jowa, No. 3621 ; Tyller, N. Y. B. G., No. 259; 
Lindheimer, Fl. texana, No. 1175. 



214 



280 



Tree; leaves on shoots 5-lobed, angular, with deliquescent teeth, deeply 
auriculate at base; leaves on trunk nearly trilobate, cuneately-decurrent 
toward base, with inconspicuous teeth; leaves on shoots broadened, 
3— 5 -lobed, the lobes weakly defined, the teeth and lobes deliquescent, 
acuminate, with broadly cordate base; heads not more than 1 or 2 (to 2.5 cm 
in diameter), the lateral sessile; fruits clavately thickened toward the 
apex, glabrous, with minute subulate style. 

Occurs in gardens of the south coast of the Crimea. Native to E. and SE 
states of the U. S. A. and Texas. Cultivated in gardens and along avenues of 
southern and central Europe and of N. America as a valuable tree species 
with ornamental and technical applications. 

A hybrid between P. o r i ent a li s L. and P. o c c id e nt al i s L., described 
as P. ac e r i f o 1 ia W., is to be found in the gardens and along the avenues 
of W. Europe, where it is known as "London" plane tree (cm. Henry a. Flood., 
The History of the London Plane. Proceed. R. Irish. Academy Vol.XXXV, sect. 
B, No. 2. Dublin, 1919). 



Family LXXVII. ROSACEAE JUSS. 

Flowers cyclic, actinomorphic in USSR species, bisexual, less often 
diclinous, solitary or in inflorescences of various types. Perianth mostly 
biseriate, in rare cases petals absent (A 1 c hi m i 1 1 a.). Sepals and petals 
usually 5, less often 3, 4, 6, 8, or many; sepals, petals and stamens often 
arranged on edge of flat, patelliform, cuplike, or goblet -shaped inflorescence 
axis —the receptacle or hypanthium (sometimes incorrectly called calyx 
tube); sepals herbaceous, mostly persistent, sometimes outer sepals present. 
Stamens mostly 2—4 times as many as sepals, or else of indeterminate 
number, occasionally only 1—5; filaments free. Carpels as many as or 
2—3 times as many as sepals or of indeterminate number, free or united 
with inner wall of hollow flower axis, unilocular, mostly with 2 pendulous 
or erect anatropous ovules, mostly separated from stamens by broad 
glandular disk. Style at apex or on ventral side, very rarely on the 
back. Fruits very varied: follicle, achene, stone-fruit, sometimes — owing 
to aggregation with expanding flower axis — forming pseudocarp or 
aggregate fruit. Seeds mostly small, usually with little developed 
endosperm. — Trees, shrubs, perennial herbs, rarely biennial or annual, 
with a spiral, very rarely opposite leaf arrangement, with stipules often 
adnate to petioles, less often exstipulate. 

Spiraea opulifolia L. — (?)— Tertiary of Balkh. (Chingistai). 

Cotoneaster vulgaris L. — interglacial Mindel-Riss of V. -Don 
(Livhvin); C.sibirica Krysht. et Bors. in Miocene of Ob' (irtysh River, 
near Tara). 

Purus communis L. — Akchagyl and Apsheron Pliocene of E. Transc. 
(S. Kakhetia, Shiraki Steppe); Pirus malus L. — interglacial Riss-Wurm 
of U. Dnp. (Murashka); Pirus sp. — Sarmatian [Upper Miocene] of Bl. 
(Krynka), in interglacial Riss-Wurm of V. -Kama (Galich). 

Sorbus torminalis L. — postglacial of E. Transc. (Pasanauri); 
Sorbis praetorminalis Krysht. et Baik. — Sarmatian of BL 
(Amvrosievka); Sorbus aucuparia L. — cave deposits of the 



215 



281 



Aurignacian, Wurms, Crimean glaciation (mountainous part of the Crimea); 
Sorb us aria Crantz. — Quaternary of Dag. (Tarku-Tau). 

Crataegus (?) furuhjelmii Heer — Tertiary of Sakh. (Due); 
C.melanocarpamaeotica Krysht.— Tertiary Maiotis deposits of 
Bess. (Seimeny); C. o xy a c ant h a: in post-Pliocene tuffs of Cisc. 
(Mashuka); C. p r a e mo n ogy na Krysht. — Sarmatian of Bl. (Krynka); 
C.fominii Krysht.— Miocene of V. -Kama. (Sterlitamak District). 



Key to Subfamilies 

1. Fruit an aggregate follicle; follicles 1 - or few-seeded, dehiscing along 
inner suture; flowers small. Shrubs, less often herbs, usually exstipulate 
Subfamily 1 . Spiraeoideae. 

+ Fruit different; stipules present 2. 

2. Herbs, sometimes shrubs, and then shoots or branches furnished with 
spicules or spines; fruit an achene, less often a stone-fruit, sometimes 
an aggregate fruit Subfamily 3. Rosoideae. 

+ Trees or shrubs; branches sometimes with spines; fruit a pseudocarp 
(pome) or drupe 3. 

3. Carpels 5, 1 or 2, united with inner side of hypanthium and often 

connate; a pseudocarp is formed at maturity 

Subfamily 2. Pomoideae. 

+ Carpel 1, not united with flower axis; fruit a drupe 

Subfamily 4. Prunoideae. 



Subfamily 1. SPIRAEOIDEAE Agardh,* Class.pl. (1825) 20; Focke in 
Nat. Pfl.III, 3 (1888) 13.- Spiraeaceae Dumort., Comm. bot. (1822) 59, 
Maxim, in A. H. P. VI, 1 (1879)163. 

Hypanthium flat, infundibular or campanulate, calycle absent; sepals 
and petals 5; stamens 15—20; pistils usually 5 (1—7); ovary with several, 
sometimes only 2, mostly pendulous ovules; fruit an aggregate follicle 
of free or more or less connate (sometimes completely united) fruitlets 
dehiscing by sutures (follicles). Shrubs, less often perennials with alternate 
simple or pinnate leaves; stipules sometimes persistent. Flowers white 
or pink. 

1. Leaves simple 3. 

+ Leaves pinnately compound 5. 

2. Flowers large, white; ovaries connate nearly to the apex; fruit 
capsulelike, of 5 follicles connate to the apex, dehiscing by sutures; 
seeds with winged margin 723. Exochorda LindL 

+ Flowers small; ovaries free or connate to not higher than the middle; 
fruit an aggregate follicle; follicles free or connate only at base; 
seeds not winged 3. 

3. Follicles much inflated, coriaceous; seeds with hard, shining testa; 

tall shrubs with large, cordate 5 (3)-lobed leaves . . . 

717. Physocarpus Maxim. 

* Treatment by A.I.Poyarkova. 



216 



282 



+ Follicles not inflated, cartilaginous; seeds with coriaceous testa; 

leaves entire or dentate, rarely shallowly trilobate at the apex .... 4. 

4. Dioeceous shrubs with small flowers in long racemes, these in turn 

in a large panicle; leaves oblong, entire, thickish 

719. Sibiraea Maxim. 

+ Flowers bisexual, in corymbiform or umbellate inflorescences or in 

terminal panicles 718. Spiraea L. 

5. Shrub with narrow linear, pinnate leaves not more than 5 mm broad, 
with 20—35 pairs of leaflets 1—2 mm long; flowers pink, in racemes, the 
latter arranged in a panicle 722. Spiraeanthus Maxim. 

+ Leaflets (l)2— 11 cm long; leaves large, lanceolate or triangular in 

outline; flowers white 6. 

6. Shrubs with pinnate leaves; flowers ca. 1 cm in diameter, in a pyramidal 
or ovoid panicle 721. Sorbaria A.Br. 

+ Perennials usually with bip innate or tripinnate leaves; flowers small, 

in spiciform racemes arranged in panicles 

720. Aruncus (L.) Adans. 



Genus 717. PHYSO CARPUS * MAXIM. 

Maxim, in A. H. P. VI ( 1879) 109. 

Shrubs with deciduous stipules and palmatilobate dentate leaves. 
Flowers bisexual, white, in simple corymbiform inflorescences at tips of 
short lateral shoots, 5 -merous; stamens 30—40; pistils 1—5, more or less 
connate, with 2—4 anatropous ovules. Follicles much inflated, coriaceous, 
dehiscing at the apex. 

1 Ovary and follicles glabrous or with sparse hairs only at the apex; 

leaves glabrous, usually with much elongated middle lobe 

*P. opulifolia (L.) Maxim. 

+ Ovary, follicles and hypanthium densely pubescent 2. 

2. Leaves grayish-tomentose below; follicles slightly longer than sepals 

1. P. amurensis Maxim. 

+ Leaves hairy only along the veins; follicles as long as or slightly 

shorter than sepals 2. P, ribesifolia Kom. 

1. P. amurensis Maxim, in A. H. P. VI (1879) 221; Kom., Fl. Mansh. U 
(1904) 453; Kom. and Alls., Opred. rast. Dal'nevost. kraya II (1932) 627. - 
Spiraea amurensis Maxim., Prim. Fl. amur. (l 859) 90. — Op u 1 a s t e r 
amur en s is O. Kuntze in Rev. Gen.pl. II (l 891 ) 949. — Ic. : Kom. and Alis., 
I.e., Plate 191, Fig. 2. 

Tall, profusely branching shrub with smooth chestnut -colored shoots 
and branches covered with brownish-gray bark peeling in strips; leaves 
cordate, with short, acute or subobtuse lobes, dark green and glabrous 
above, grayish below with a stellate pubescence; inflorescence 
5— 25 -flowered, with tomentose axis, pedicels, and hypanthium; flowers 
16— 18mm in diameter; petals oblong, white, hairy outside; ovaries 2—4, 
connate at base; follicles with mostly 3, sometimes 2 or 4 seeds. Fl. May, 
fr. August. (Plate XVII, Figure 1 ). 

• From the Greek phy sa , Dladder and c arpos, fruit. 



217 



283 



284 



Stony slopes, shrub thickets, mixed forests, often on limestones. — 
Far East: Ze.-Bu. — Bureya Range, Uss. Gen.distr.: Jap.-Ch.: Manchuria 
and N.Korea. Described from the Bureya Mountains. Type in 
Leningrad. 

Economic importance. Cultivated as an ornamental plant. 

2. P. ribesfolia Kom. in Bull. Jard. Bot. Acad. Sc. URSS, XXX (1932) 
202; Kom. and Alis., Opred. rast. Dal'nevost. kraya II (1932) 627.- (?) 
Opulaster amurensis Nakai in Fl. sylv. Kor. IV (1916) 29 (non 
O. Kuntze). — P. amurensis var. concolor Kom. in sched. 

Shrub, profusely branching, with dense crown and ramentaceous dingy 
gray bark; leaves 3—5 -pinnatilobate, with acute lobes and cordate base, 
glabrous, with simple and stellate hairs only along the veins below; 
pedicels, hypanthium, sepals, ovary and follicles glabrous, the latter slightly 
shorter than or as long as sepals. Fl. May, fr. August. 

Stony slopes.— Far East: Ze. -Bu. (Bureya Range), Uss. Gen.distr.: 
Jap.-Ch., N.Korea. Described from the vicinity of Vladivostok. Type 
in Leningrad. 

Economic importance. Ornamental plants. 

• P. opulifolia (L.) Maxim, in A. H. P. VI (1879) 220. - Spiraea opuli - 
folia L., Sp.pl. (1753) 489.- Neilla opulifolia Benth. et Hook., 
Gen. pi. I (1865) 612. — Op u 1 a s t e r opulifolius O. Kuntze, Rev. gen. pi. 
II (1891) 9490.- Ic: Rchb., Ic. Fl. Germ. XXIV (1909) tab. 153. 

Tall shrub, to 3 cm high, with somewhat nodding branches and glabrous 
very light brown shoots; leaves 3— 5 -lobed, the middle lobe usually 
prolonged, glabrous; inflorescences glabrous or rather sparsely pubescent; 
flowers 10— 12 mm in diameter; stamens 30; pistils 3—5; follicles 2 or more 
times as long as sepals, glabrous or sparsely pubescent at the apex. 

Growing wild in NE America, from Quebec to Georgia; in the USSR 
firmly established in cultivation as a hardy ornamental shrub. 



Genus 718. SPIRAEA* L. 

L.Sp.pl.(1753) 489. 

Flowers bisexual, in corymbiform, umbellate, or paniculate inflorescences, 
with 15—30 stamens and 5 — (3— 8) pistils alternating with sepals; ovaries 
free, with several anatropous pendulous ovules. Shrubs with simple leaves, 
more or less dentate, less often entire or lobate at the apex; exstipulate. 

Economic importance. Nearly all species are ornamental nectariferous 

1. Flowers in panicles 2. 

+ Flowers in corymbiform or umbellate inflorescences 4. 

2. Panicles broadly ovoid, very sparse, with very remote branches; 
flowers white 4. S. baldschuanica B. Fedtsch. 

+ Panicles dense, ovoid, ovoid -cylindrical or pyramidal; flowers bright 

pink 3. 



From the Greek speiraia, a shrub name used by Theophrastus. 



218 



3. Young shoots, petioles, inflorescence axis, and hypanthium densely 
covered with ferruginous tomentose pubescence; leaves broadly 
elliptic or ovate (1.5—2.5 times as long as broad), dentate usually 

from above the middle 2. S. humilis A. Pojark. 

+ Young shoots glabrous or with rather sparse light pubescence; 
inflorescence axis and pedicels pubescent; hypanthium sparsely 
hairy; leaves long-elliptic or elliptic -lanceolate (3—4 times as long 
as broad), dentate almost from base 1. S. salicifolia L. 

4. Flowers in compound corymbiform inflorescences 5. 

+ Flowers in simple corymbs or umbels 6. 

5. Inflorescence 2.5 — 9 cm in diameter (mostly 5—6 cm), with long, 
profusely branching lower branches; pedicels glabrous or slightly 
pubescent; leaves 2.2— 7.5 cm long (usually ca. 4 cm), dentate almost 
from base 3. S. betulifolia Fall. 

+ Inflorescence 1.5—3 cm in diameter (mostly 2 cm), with little- 
protruding lower branches; pedicels gray with dense pubescence; 

leaves 1—3 cm long, often dentate only at the apex or entire 

4. S. beauverdiana C. K. Schn. 

6. Flowers in axillary sessile umbels with a leaf rosette at the 

base 7. 

+ Inflorescences terminating axillary leafy shoots 8. 

7. Leaves of sterile shoots — except the very lowest — broadly cuneate, 

with 3 large lobelike teeth at the apex; pedicels 2—6 mm long 

22. S. aquilegifolia Pall. 

+ Leaves of sterile shoots oblong, entire or with 2—5 small teeth at the 

apex; pedicels 5 — 15 mm long 21. S. hypericifolia L. 

8. Style arising from ventral side of follicle 9. 

+ Style arising from dorsal side of follicle 12. 

9. Fruiting sepals erect; buds ovate, with 6—8 free exterior scales; 

petioles several times longer than buds 9. S. elegans A. Pojark. 

Fruiting sepals recurved; buds oblong, flattened, more or less equal 

to petioles, with 2 sharp -pointed exterior scales 10. 

10. Leaves oblong-elliptic, with simple acute teeth at the apex or from 

the middle; flowering shoots 1.5— 6 cm long 7. S. flexuosa Fisch. 

+ Leaves ovate, duplicato -dentate from base or from the middle .... 11. 

11. Shoots light brown, slender; leaves broadly ovate with rounded or 
truncate base, the secondary teeth equal, small; flowering branchlets 

2—5 cm long; inflorescences subumbellate 

8. S. ussuriensis A. Pojark. 

+ Shoots light yellow or brownish yellow, thicker; leaves often cuneate 
at base, the secondary teeth unequal; flowering shoots 3—14 cm 
long; inflorescence corymbiform, sometimes with very scattered 
lower flowers 6. S. chamaedryfolia L. 

12. Leaves with long, appressed sericeous pubescence on both sides or 
only below; stamens longer than petals; fruiting sepals recurved; 
leaves with few teeth at the apex U.S. sericea Turcz. 

+ Pubescence absent or different 13. 

13. Leaves with a whitish bloom below; inflorescences small, very dense; 
flowering shoots 1 — 3 cm long 14. 

+ Leaves green below, without bloom 15. 



219 



286 



14. Leaves oblanceolate or linear -lanceolate, mucronate, dark green 
above; branches usually long, virgate, with numerous flowering 

shoots 18. S. alpina Pall. 

+ Leaves oblong-oboval, with obtuse or rounded apex, usually glaucous; 

branches short, with 1—5 flowering shoots 

14. S. tianschanica A. Pojark. 

15. Stamens shorter than petals; fruiting sepals erect; leaves with large 
teeth at the apex or shallowly trilobate 16. 

-+ Stamens longer than petals; leaves entire, those on sterile shoots 

dentate, or else with few small teeth confined to the apex .... 18. 

16. Quite glabrous shrub with broad leaves 15. S. trilobata L. 

+ Shrubs with pubescent leaves and shoots, often other parts also 

pubescent, or leaves narrow, lanceolate 17. 

17. Hypanthium, sepals, and pedicels glabrous; bracts absent. Shrubs 

2— 4 m high 17. S.pubescens Turcz. 

+ Hypanthium and sepals pubescent; pedicels with linear bract above 

the middle. Shrubs 30— 70 cm high 16. S. pilosa Franch. 

18. Fruiting sepals recurved; pedicels 6 — 12 mm long; buds with several 
free exterior scales 19. 

+ Fruiting sepals erect; pedicels 2— 5 mm long (to 8 mm in fruit); buds 

covered by mitra of connate scales 20. 

19. Leaves narrow, elongate -elliptic or linear -lanceolate; shoots 
slender, virgate; entire plant glabrous; flowering shoots 2—3 cm 

long; flowers 6— 7 mm in diameter . . 12. S. dahurica Maxim. 

+ Leaves broader; shoots thicker, strong; flowering shoots 3— 8 cm 

long; flowers 7— 9 mm in diameter 10. S. media Schmidt. 

20. Leaves on sterile shoots with 3 parallel longitudinal veins, regularly 

dentate or crenate from the middle or nearly from base 

18. S. crenata L. 

+ Leaves on sterile shoots with one midrib, entire or with 2 or 3 teeth 

at the apex 21 . 

21. Flowers 7— 10 mm in diameter; pedicels 2— 5 mm long, to 8 mm long 

in fruit; leaves on sterile shoots 12—22 mm long 

19. S. lasiocarpa Kar. et Kir. 

+ Flowers 10— 12 mm in diameter; pedicels 8— 18 mm long, those of 
uppermost flowers 4 mm long; leaves on sterile shoots 20— 35mm 
long 20. S. ferganensis A. Pojark. 



Subgenus 1. PR0T0SPIR4EA. Nakai, Fl. sylv. Koreana IV (1916) 12.- 
Inflorescences developing at ends of new annotinous long leafy shoots, arising 
from base of shrub or from old, rarely from second-year branches. 

Section 1. SPIRARIA Ser. in DC, Prodr. II (1825) 514, (ex parte); 
C.K. Schn., III. Handb. Laubh. I (1906) 480.— Inflorescence an oblong, more 
or less pyramidal ovoid -cylindrical panicle. 

Small number of species, all — apart from the two USSR species — 
distributed in N. America. 



220 



1. S. salicifolia L., Sp.pL (1753) 489; Pall., Fl. Ross. I (1784) 36 

(var. major et alpestris); Turcz., Fl. baic. -dah. I, 361 ; Ldb., Fl. Ross. II, 
1,15; Shmal'g., Fl. I. 314; Kom., Fl. Mansh. II (l 904) 454; Ej., Fl. kamtsch. II 
(1929)229; Hulten, Fl. Kamtch. Ill (1929)427; Kom. and Alis., Opred. rast. 
Dal'nevost. kraya II (1932) 627; Kryl., Fl. Zap. Sib. VII, 1457.- S. salic i - 
folia a Ian ceo lata et j3 alpestris Maxim, in A. H. P. VI (1879) 210.- 
Ic: Pall.. I.e., t. XXI, XXII.- Exs.: HFR.No. 126. 

Shrub 1— 2 m high, with smooth, usually glabrous shoots; leaves 4 — 10 em 
long, 1.5—4 cm broad, elongate -lanceolate or long-elliptic, acute, cuneately 
tapering toward base into short petioles (2 — 7 mm), glabrous or pubescent 
below along the midrib, with very prominent veins, serrate nearly to base; 
panicles ovoid -cylindrical or pyramidal, usually 5—12 cm long, sometimes 
rounded -ovoid and 1.5—2 cm long, the specimens with such impoverished 
inflorescences usually also with smaller leaves (f. alpestris Pall.) 
and occurring mainly in the northern part of the distribution area; 
inflorescence axes and short pedicels usually more or less covered 
with yellowish pubescence, rarely quite glabrous; flowers 8—12 mm in 
diameter, pink, the stamens twice as long as petals; follicles smooth, 
with style arising from the dorsal side almost at the apex, the upper parts 
recurved. Fl. June— August, fr. July— September. 

Riverbanks, flooded meadows, tussocks of forest and meadow bogs; 
nearly always forming thickets, low -growing in open places and attaining 
2m in shadier sites.— Arctic: An.; W.Siberia: Ob — Ob River basin 
between 56.6 and 61° N., U. Tob. (very rarely on the Tobol River and in the 
Naryni River valley); E. Siberia: Yenis. (to the latitude of the Lower 
Tunguska River), Lena-Kol. (as far north as Verkhoyansk; known from the 
Indigirka and Kolyma rivers), Ang.-Say. (apparently with the exception 
of the SW part), Dau.; Far East: Kamch., Ze.-Bu., Uda, Uss., Sakh. 
Gen. distr. : Centr. Eur.: from Carniola, Upper Austria and Bohemia to 
Lithuania, Volhynia, and Bukovina; Mong. (Tuva ASSR, N. Mong.); Jap.-Ch. 
(Manchuria, N. Korea, Japan); reported for Alaska and Sitka ? Described 
from Siberia. Type in London. 

Economic importance. Long and well established in horticulture, 
especially the large -flowered variety with pale pink flowers — 
var. g r an d i f 1 o r a C. Koch, usually known as S. gr an d i fl o r a Lodd. 
Replaced in America by the closely related species S. alba Du Roi and 
S.latifolia Borkh., which are quite often cultivated in the gardens 
and parks of the Soviet Union. 

2. S. humilis A. Pojark. in Addenda VIII, p. 375. 

Shrub 25— 50 cm high; young shoots densely covered with ferruginous 
tomentose pubescence; leaves 2.5—6.5 cm long, 1—3.3 cm broad, elliptic, 
less often ovate, rather broad (1.5—2.5, rarely 3 times longer than broader), 
with cuneate, less often rounded base, acute, glabrous above, with rather 
sparse sericeous hairs along the main veins below, usually with acute, 
mostly simple teeth only from the middle or only near the apex, rarely 
teeth present also in lower part of leaf; lower leaves on shoots sometimes 
entire; petioles 2— 4 mm long, ferruginous -tomentose; flowers bright pink, 
in compact, broad, ovoid or pyramidal -ovoid panicles, 2.5 — 10 cm long, 
2—6.5 cm in diameter, with dense ferruginous -tomentose pubescence on 



221 



289 



axes, pedicels, and hypanthium at least in its lower part; pedicels 
1.5 —3 mm long, thick; sepals triangular, acute; stamens much longer 
than petals. Fruits unknown. July. 

Mossy tussocks in boggy deciduous forests.— E. Siberia: Lena-Kol.; 
Far East: Uda, Uss., Sakh. Endemic ? Described from Udinskoe on the 
Amgun River. Type in Leningrad. 



Section 2. CALOSPIRA C.Koch in Gartenflora III (1854) 397; C.K.Schn. 
Handb. Laubholz. I (1906) 467. — Flowers in broadly corymbiform or ovoid 
panicles. 



Cycle 1 . Betulifoliae A. Pojark. — Flowers white or pale pink, in 
dense corymbiform panicles, the stamens twice as long as petals; 
fruiting sepals recurved; follicles linear -lanceolate, with truncate apex; 
style arising from dorsal side. 

3. S.betulifolia Pall., Fl. Ross. I (1784) 33; Ldb., FL Ross. II, 14; Kom.., 
Fl.Mansh. II (1904) 456; Kom., Fl. Kamtsch. II (1929) 230; Kom. and Alis.. 
Opred. rast. Dal'nevost. kraya II (l 932) 627. — S.betulae folia var. 
typica Maxim, in A.H. P., VI (1879) 208 (fl.ochroleuco et fl.roseo).- 
S.beauverdiana Hulten. Fl. kamtch. Ill (192 9) 38 (ex parte) non 
C.K. Schn. - Ic: Pall., 1. c, tab. 16. 

Shrub ca. 50—60 cm high, with dense globose crown and very leafy glabrous 
and pubescent shoots covered — like the old branches — with light brown 
bark; leaves elliptic or oboval with cuneate base, less often broadly ovate 
with rounded base, short -petioled, 2 .2— 7.5 cm long. 1—5 cm broad, mostly 
ca. 4 cm long, 2.5 cm broad, light below, prominently veined, glabrous on 
both sides or pubescent along the veins, crenate-dentate on the margin 
mostly almost from base; inflorescence 2.5— 9 cm in diameter (usually 
5—6 cm), often with more than 100 flowers, the lower lateral axis much 
branched, to 9 cm long; pedicels and inflorescence axis glabrous, less 
often sparsely pubescent; flowers 7— 9 mm in diameter, white or pinkish 
(drying slightly yellowish, which may explain Maximowicz's description 
off. ochroleuc a); follicles 3—4 mm long, pubescent or glabrous. Fl. from 
end of June to beginning of September, fr. from July. (Plate XVII, Figure 3). 

Dry sparsely wooded and open mountain slopes, rock streams, peat bogs, 
mainly deciduous and Siberian stone pine forests; single shrubs or in small 
groups.— E.Siberia: Lena-Kol. (SE part); Far East: Kamch. (together with 
S.beauverdiana C.K. Schn. but not very often, and apparently rarely 
pure, mostly as a hybrid with S.beauverdiana), Okh. (Taui Bay), Ze. -Bu. 
(as far west as Selendzha River, a tributary of the Zeya), Uda, including 
the Shantar Islands. Uss., Sakh. Gen. distr. : Jap.-Ch. (Chihli Province), 
Jap. Described from the Maya River, a tributary of the Aldan. Type in 
London. 

Note. Usually, D. 1 u c i d a Dougl. from NW America (from British 
Columbia to Oregon) closely resembling S.betulifolia in its habitus, 
is indicated as being very closely related to it. This species, however, 
is distinguished from C.betulifolia by such essential characters as 



222 



erect fruiting sepals and by follicles tapering to the apex, their style 
arising very near to the ventral side, which precludes such a close 
relationship. Hulten (l. c.) put forward the hypothesis that Pallas's 
name S.betulifolia Pall, refers precisely to this N. American 
species and not to the E. Asian one. This assumption originated from 
his mistaking the specimen in the BIN Herbarium for Pallas's authentic 
specimen: it bears a close resemblance in leaf arrangement and shape of 
inflorescence to the description of Steller's specimen in Fl.Ross., 
Plate 16, from which S. b etu 1 if o 1 ia Pall, has been described. The 
specimen of S. lucid a referred to bears the label "betulaefolia" in 
Fisher's handwriting (this plant clearly belongs to his herbarium); 
below there is a note in Regel's writing: "Herb. Pallas," which must 
have caused Hulten's error. 

Economic importance. The low -growing, abundantly flowering shrubs 
of this species are used as ornamental plants, especially for border settings. 

S.betulifolia Pall. X b e au v e r d i an a C. K. Schn. — these hybrids 
are rather frequent in the northern part of the S.betulifolia distribution 
area, in Kamchatka, the Okhotsk coast, and Sakhalin, but occur rather rarely 
in the southern parts of its distribution area. In their characters they 
approximate one or other of the parent species. 

4. S.beauverdiana C.K. Schn.in Bull. Herb. Boiss.,2 ser., V (1905) 348; 
111. Handb. Laubh. I (1906) 478 (van typica et var. S t e v e ni); Kom., Fl. 
Kamtsch. II (1929) 231; Hulten, Fl. kamtch. Ill (1929) 38 (ex parte). - 
S.chamaedryfolia (non L.) Cham, et Schlecht. in Linnaea II (1827) 2. — 
S. crenata Mertens in Linnaea V (1830) 63, non L. — S. betulaefolia 
var. nan e 11a Kom., 1. c, 231 . - S. s t e v e n i Rydb. in N. Amer. Fl. XXII, 3 
(1908) 247.; 

Small, profusely branching shrub 15—30 cm high, the shoots slender, 
ribbed, pubescent when young, later glabrous; leaves short -petioled, 
1—3 cm long, 0.5— 1 .5 cm broad, mostly 2 : 1 .2 cm, elliptic or oval, often 
elongated (3 : 1 ), obtuse, usually rounded, rarely cuneate at base, glabrous, 
light below, with very prominent veins, crenate- or serrate -dentate, 
often only in upper part, rarely entire; inflorescences small, 1.5 —3 cm 
in diameter (mostly 2 cm), very dense, the lower branches to 1.5 cm 
long, little -protruding; axes short (to 3 cm long); pedicels grayish with 
dense velutinous pubescence; flowers 4—6 (7) mm in diameter; follicles 
2.5—3 mm long, mostly pubescent. Fl. June, fr. from end of July. 
(Plate XVII, Figure 2). 

Habitats as for the preceding species.— Arctic: Chuk., An.; E.Siberia: 
Lena-Kol. (SW part); Far East: Kamch., Okh., Ze.-Bu. (upper part of Zeya 
and Bureya river basins), Uda, Uss., very rarely, known only from the 
BotchiRiver,Sakh. Gen. distr. : Jap.-Ch.: N. Japan, Kurile Islands and 
Ber., NW tip of America (Alaska to Yukon). Described from Japan. Type 
in Vienna. 



Series 1. Decumbentes A. Pojark. — Flowers white, in very sparse, 
sometimes oblong panicles; stamens half as long as petals; fruiting sepals 
erect; styles arising from dorsal side very near the apex. Apart from 



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our species, S.decumbens Koch, and S.haqueti Fenzl. et Koch., from 
the limestones of Italy and Carinthia, also belong to this series. 

5. S. baldschuanica B. Fedtsch. in Consp. fl. turk. Ill (1909) 6. 

Low, branching shrub with glabrous, reddish-brown shoots; leaves firm, 
glaucous -greenish, lighter below, glabrous, oboval, broadly elliptic, or 
subrhomboid, cuneately tapering to base, with a short (1.5—3 mm) petiole, 
acute, less often obtuse, entire or unequally dentate above the middle, 
(5)l0— 22 mm long, (2.5)4.5—15 mm broad; panicles to 8 cm in diameter, 
to 12 cm long, very loose, with very remote, slender branches usually 
3—5 cm, sometimes to 9 cm long, bearing at the base one bract, which 
is sometimes displaced nearly to middle of axis; pedicels slender, filiform, 
with a lanceolate bracteole above the middle; flowers white, 7 mm in 
diameter, the sepals and hypanthium pubescent outside; follicles pubescent, 
3 times as long as sepals. June. 

Limestones on mountain slopes.— Centr. Asia: Pam.-Al. — known only 
from southern slope of the Gissar Range (from the Varzob River), from 
the Gazimailik Range and the Vakhsh Range (Mts. Changlak and Sevistan). 
Endemic. Described from the Sevistan Mountains. Type in Leningrad. 

Economic importance. May be used as ornamental plants. 



Subgenus 2. METASPIRAEA Nakai, Fl. sylv. Koreana IV (1916) 12.- 
Inflorescences developing from buds of preceding year's shoots, corymbiform 
or umbellate, at tips of short leafy branchlets or sessile. 



Section 1. CHAMAEDRYOM Ser. in DC, Prodr.II (1825) 542.- 
Flowers in simple corymbiform or umbellate inflorescences terminating 
leafy lateral shoots, less often in sessile umbels with leaf rosette at base. 



Series 1. Chamaedryfoliae A. Pojark. — Inflorescences terminating leafy 
shoots; stamens twice as long as petals; fruiting sepals recurved; style 
arising from ventral side of follicle; shoots ribbed; buds more or less 
equal to petioles, flattened, oblong, with 2 sharp -pointed exterior scales.— 
Three species occur in the Soviet Union. 

6. S. chamaedry folia L., Sp.pl. I (1753) 489; Ldb., Fl. Ross. II (1844) 4 
(ex parte); Bge. in Ldb., Fl. Alt. II (1930) 213; Kryl., Fl. Zap. Sib. VII, 
1454.- S.ulmifolia Scop., Fl. carniol. I (1772) 349. - S. chamaedry - 
folia var.ulmifolia Maxim, in A. H. P. VI (1879) 186 (ex parte). - 
Ic: Pall., Fl. Ross. I (1784) tab. 15 (except for lowest figure on left of 
plate). 

Shrub 80—150 cm high, with glabrous, mostly erect shoots, from pale 
yellow to very light brown; leaves with short (2—5 mm) petioles, glabrous, 
or with beards below at base, broadly ovate or oblong -ovate, acute, with 
rounded or cuneate base, unequally dentate or duplicato -dentate nearly from 
base or from above the middle; leaves on sterile shoots sometimes incised- 
dentate, those on flowering shoots often with slightly developed, subobtuse 



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292 



teeth or entire, 1.5—6 (lO)cm long, 1— 4 cm broad; flowering shoots 3— 14cm 
long, only the uppermost sometimes 2cm long; inflorescences 6— 20 -flowered, 
corymbiform, often with remote lower pedicels, the latter glabrous, 
13— 20 mm long; corolla 10— 15mm in diameter; follicles glabrous or 
appressed -hairy. Fl. June, fr. July. (Plate XVIII, Figure 9). 

Open forests, forest edges, banks of small rivers, shady slopes, also 
stony slopes, sometimes forest meadows and subalpine meadows.— 
W. Siberia: Ob — SE part between the Ob and Yenisei rivers, Irt. — SE part, 
Alt.; E. Siberia: Ang.-Say. — only in area adjacent to the Yenisei; 
Centr. Asia: Balkh. — only in the far SE part (near Zaisan), Dzu.-Tarb. 
(Saur, Tarbagatai, and the Dzungarian Ala-Tau). Gen. distr.: Centr. Eur. — 
from Carniola, Austria, and Hungary, to the northern part of the 
Balkan Peninsula and the Carpathians; Mong. — Tuva ASSR. Described 
from Siberia. Type in London. 

Economic importance. Widely used as ornamental plants. Nectariferous. 

7. S. flexuosa Fisch. ex Cambess. in Ann. sc. nat. I (1824) 365; Ser. in 
DC, Prodr.II (1825) 542; Turcz., Fl. baic.-dah. I, 357; Ldb., Fl. Ross. II, 

1, 13; Kom., Fl. Mansh. II (1904) 458 (ex parte); Kom. and Alis., Opred. rast. 
Dal'nevost. kraya II (l 932) 62 7 (ex parte). — S.chamaedrifolia var. 
flexuosa Maxim, in A. H. P. VI (l 879) 1 86. - Ic. : Cambess., I.e., tab. 26. 

Shrub to 1.5 m high, with slender, sometimes geniculate shoots, pale 
yellow or brownish yellow; leaves oblong -elliptic or oblong -oval, rarely 
broader, acute, with cuneate, less often rounded base, with remote, acute, 
sometimes inconspicuous teeth diminishing upward from middle of blade 
or higher, quite glabrous or else very sparsely hairy below; flowering 
branches 1.5—6 (8) cm long, 4— 10 -flowered; pedicels glabrous, 0.7— 1 8 mm 
long; corolla 8— 10 (l 2) mm in diameter, sometimes pinkish; follicles 
usually appressed -hairy at the apex. Fl. June, fr. August. (Plate XVII, 
Figure 8). 

Forest edges and often deciduous -and -pine and mixed forests, in forest 
meadows, sometimes subalpine meadows, on banks of small rivers, less 
often on slopes among shrub thickets.— W. Siberia: Ob (easternmost 
part only), Alt. — on the Ang. -Say. border; E.Siberia: Lena-Kol. —known 
only fromthe Maya River, Ang.-Say., and Dau. (W. ); Far East: Uss. (rarely). 
Gen. distr.: Mong. — Tuva, mountainous part of N. and Centr. Mong., 
Jap. -Ch. — N.Korea (Segelsu River valley). 

Described after a cultivated specimen. Type in Geneva. 

Economic importance. Has long been cultivated in European gardens, 
but of rare occurrence. Hardly cultivated at all in the Soviet Union. 

8. S. ussuriensis A. Pojark. sp. n. in Addenda VIII, p. 375 — S. chamae- 
dryfolia Maxim., Prim. fl. amur. (1859) 90 (non L.); Rgl. in Mem. Acad. 
Sc.St.-Petersb., VII, ser., IV, 4 (1861) 52; Kom., Fl. Mansh. II (1904) 457; 
Kom. and Alis., Opred. rast. Dal'nevost. kraya II (1932) 6 97.— S. flexuosa 
var. 1 at i f o 1 i a Fisch. ex Maxim., 1. c, 91 (ex parte et pro synon.). — 

S. ulmifolia non Scop. ) Nakai, Fl. sylv. Koreana IV (1915) 20.— Ic . : Kom. 
and Alis., 1. c, Plate 192, Figure 4; Nakai, 1. c, Plate 8. 

Shrub ca. 1 m high, with slender, often flexuous light brown shoots; 
leaves slender, broadly oval, rounded -cuneate or truncate at base, acute 
or subobtuse, glabrous above, with hairs along the midrib below and with 



225 



293 



beards in the angles, duplicato -dentate, the secondary teeth more or less 
equal, small, acute; flowering branches 2—5 cm long, with 3—5 leaves not 
differing in shape of dentation from those on vegetative shoots; flowers 
9— 15mm in diameter, white, in umbellate 4— 12 -flowered inflorescences 
(pedicels not remote as in S.chamaedryfolia); pedicels 7—15 mm long, 
glabrous like the calyx, often with glaucous bloom; follicles usually 
appressed -hairy only in upper part. Fl. June, fr. July. (Plate XVII, 
Figure 10). 

Slopes and rocks, both in mixed forests and in open spaces.— Far East: 
Ze.-Bu. (only Bureya Range), Uda (southernmost part), Uss. Gen.distr.: 
Jap.-Ch. — Manchuria and N.Korea. Described from the Khekhtsyr Range 
in the Ussuri area. Type in Leningrad. 

Economic importance. Ornamental and nectariferous. 



Series 2. Elegantes A. Pojark. — Flowers in few -flowered corymbs, 
terminating leafy shoots; stamens twice as long as petals; fruiting sepals 
erect; style arising from ventral side of follicle; shoots ribbed; buds 
ovate, with 6—8 free exterior scales, the petioles several times 
longer than buds. — One species. 

9. S. elegans A. Pojark. sp. n. in Addenda VIII, p. 376.— S.chamaedry- 
folia Fr. Schmidt in Mem. Acad. Sc. St. Petersb., VII ser., XII, 2 (1868) 
38 (non L. ). — S.chamaedryfolia var. ulmifolia ex parte) et var . 
f lexuosa Korsh. in A. H. P. XII (1892) 333. 

Shrub with slender, slightly ribbed, often flexuous, glabrous shoots 
covered with light brown or reddish -brown bark, and with brownish-gray 
strongly ramentaceous older, slender branches; leaves 11—35 mm long, 
6—15 mm broad, on sterile shoots to 55 mm long, 26 mm broad, leaves 
with petioles 2—11 mm long, glabrous above, with beards in angles of main 
veins below, oblong-elliptic or lanceolate -oval, subacute, usually with 
cuneate base, unequally dentate and partly duplicato -dentate above base 
or at least from the middle (on flowering shoots sometimes only at the 
apex), sometimes incised -dentate on sterile shoots; flowering branches 
1—5 (7) cm long (one plant was found with sessile inflorescences); 
inflorescences 6— 14 -flowered; pedicels 7—12 mm long, to 16 mm long 
in fruit, glabrous; corolla 10— 15 mm in diameter; follicles with appressed 
yellow pubescence throughout or confined to the apex. Fl. June, fr. August. 
(Plate XVII, Figure 7). 

Stony slopes, rock streams and rocks, mostly woodless.— E. Siberia: 
Dau. (E.); Far East: Ze.-Bu., Uss. Gen.distr.: Mong. — in part of 
N. Mongolia adjacent to Dauria. Described from the village of Pokrovka 
on the Amur. Type in Leningrad. 

Note. S. elegansXussuriensis has buds like those of S. e 1 e g a n s, 
but in leaf shape it is closer sometimes to one species, sometimes to the 
other. In available specimens, sepals are recurved as inS.ussuriensis. 
Apparently not a very rare species; collected in the Bureya Range and on 
the coast (Uss.). Resembles in habitus S. u s s ur i e ns i s A. Pojark., 
from which it is distinguished by erect fruiting sepals, entirely different 
bud structure, shape of leaves and of leaf teeth. 



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294 



Cycle Mediae A. Pojark. — Flowers in loose corymbiform inflorescences 
terminating leafy shoots; stamens longer than petals; fruiting sepals divergent; 
style arising from dorsal side of follicles; shoots smooth, not ribbed; 
buds with 6—8 free scales. — Apart from the 3 USSR species, S. prostrata 
Maxim, from central China also belongs in this cycle. 



10. S. media Schmidt, Oesterr. Baumz. I (1792) 53; Shmal'g., Fl. I, 315; 
Kom., Fl. Kamtsch. II (1929)232; Kom. and Alis., Opred. rast. Dal'nevost. 
kraya II (1932) 627 (ex parte); Kryl., Fl. Zap. Sib. VII, 1455. - S. oblongi - 
folia W. et K., Ic. pi. rar. Hung. Ill (1812) 261.- S.chamaedryfolia Ldb., 
Fl. Ross. II, 14 (ex parte, non L.); Turcz., Fl. baic. -dah. I, 358. — 

S. saxatilis Turcz., I.e., 318 (nomen) et in sched. — S.confusa Rgl. 
et Koern. in Ind. sem. hort. Petrop. (1857) 57. — S. c onf us a a sub glabra 
Rgl. in Mem. Acad. Sc. St. Petersb. VII Se'r., IV, 4 (l 861 ) 52. - S. b an at i c a 
Janka in Acad. Kozl.XII (1876) 166.— S.polonica Blocki in Deutsch. bot. 
Monatsschr. X (1892) 110.- Ic: Schmidt., 1. c, tab. 54; W. etK.,1. c, 
tab. 235; Kom. and Alis., I.e., Plate 192, Fig. 3.- Exs.: HFR, No. 1215a, 
1215b. 

Shrub 1— 2 m high, with glabrous shoots pubescent when young, covered 
with light brown bark, later ramentaceous; leaves elliptic, oblong to 
sublanceolate, entire on flowering shoots, upper leaves on sterile shoots 
with few unequal teeth at the apex, thin, glabrous on both sides or with 
sparse hairs below, ciliate-margined; flowering branchlets 3— 8 mm long; 
pedicels 7—22 mm long; corolla 7— 9 mm in diameter; follicles glabrous or 
hairy, the style arising from dorsal side, slightly below the apex. Fl. from 
end of May — July (in various regions), fr. July —August. 

Undergrowth of dry forests, forest edges, open, shady and stony slopes, 
among shrubs, valley grass thickets; mostly in groups.— European part: 
Dv.-Pech., V.-Kama, U. Dnp. (SW). W.Siberia: Ob (S. part), Irt., Alt.; 
E.Siberia: Yenis. (S. part), Lena -Kol. (except N.), Ang. -Say., Dau. ; 
Far East: Kamch., Okh., Ze. -Bu., Uda, Uss. (rarely); Centr.Asia: Dzu.-Tarb. 
(in N. part — Saur). Gen. distr.: Centr.Eur.: Poland, Hungary, 
Carniola; Mong.— Tuva, N. Mong., Jap. -Ch. — Manchuria, N. Korea. Described 
Described from the Irtysh River in Siberia. 

Economic importance. One of the most common ornamental shrubs 
in park and gardens of the USSR; used mainly for hedgerows, being tolerant 
?q7 of clipping. Nectariferous. 

Note. S. mediaXcrenata = S.pikoviensis Bess.; hybrids 
showing intermediate characters between the two species are easily 
recognized; the leaves on the sterile shoots are similar to those of 
S. m e d i a in size, shape, and dentation, and to those of S. c r e n a t a in 
the presence of 3 longitudinal veins. 

11. S. sericea Turcz., Fl. baic. -dah. I (1842) 358; Maxim., Prim. fl. amur. 
(1859) 9.1. — S. c onf us a var. sericea Rgl. in Mem. Acad. Sc. St. Petersb., 
VII ser., IV, 4 (1861) 53.- S. media var. sericea C. K. Schn., 111. Handb. 
Laubh. I (1906) 456. — S. med ia Kom. and Alis., Opred. rast. Dal'nevost. 
kraya II (1932) 627 (ex parte). 

Shrub 70—180 cm high, with light brown, puberulous or tomentose shoots 
and reddish -gray branches with strongly ramentaceous bark; leaves 1—3 cm 
long (to 4.5 cm long on sterile shoots), 0.7—2 cm broad, firm, prominently 



227 



[2 95) 




PLATE XVII. 1 — Physocarpus amurensis Maxim., a) fruit; 2 — Spiraea beauverdiana C.K. 
Schn. ; 3 - S. be tul i fol ia Pall., leaf; 4 - S. p i 1 osa Franch, a) fruit; 5 - S. t r i loba ta L., leaf; 
6 - S.alpina Pall.; 7 - S.elegans A. Pojark., a) fruit, b) bud; 8 — S.flexudsa Fisch., a) fruit, 
b) bud; 9— S.chamaedrifolia L., a) leaf; 10— S.ussuriensis A. Pojark., leaf; 11 — S.dahurica 
Maxim., a) sterile shoot, b) fruit. 



228 



298 



veined, with long sericeous pubescence, often continuous and long-persistent 
below but disappearing with growth above, oval or elliptic, varying 
greatly in width (from broadly ovate to lanceolate), mostly acute, entire, 
usually with few teeth at the apex only on sterile shoots; petioles short, 
1— 2(3)mm; flowering branchlets 2— 6 cm long; inflorescences many- 
flowered; pedicels 7— 9 mm long; follicles mostly pubescent, the style 
often arising from the dorsal side very near the apex. Fl. June, fr. July. 

Usually on open, stony slopes, shrub thickets, forest edges, dry forests, 
taluses in forests; sometimes forming thickets.— E.Siberia: Dau. 
(E. part); Far East: Ze.-Bu., Uda, Uss. Gen.distr.: Jap. -Ch. (Manchuria) 
and N. Mong. Described from the Argun River in Dauria. Type in 
Leningrad. 

Note. This species is usually regarded as a variety of S. m e d i a Schmidt. 
However, S. sericea Turcz. is distinguished by very constant characters 
(persisting in cultivation) and has a specific distribution area coinciding 
with only a relatively small part of the distribution area of S. m e d i a. 
In places where both species grow together, S. sericea is encountered 
much more often than S. media (especially in the Ussuri region) and is 
confined to more open and drier habitats. 

Hybrids of S. sericea X S. media occur seldom and have intermediate 
characters. 

Economic importance. Should be cultivated as an ornamental plant. 

12. S. dahurica Maxim, in A. H. P. VI (1879) 190.- S. alp in a var. 
dahur ic a Rupr. in Bull, phys.-math. Acad. Sc. St. -Petersb. XV (1857) 362. - 
S. -alp in a Korsh. in A. H. P. XII (1892) 333 (non Pall.). 

Glabrous, spreading shrub 1—1.5 m high, with slender, virgate, often 
arched -upcurved dark brown shoots; leaves 8—20 cm long (to 30 mm on 
sterile shoots), 2—3.5 mm broad, elongate-elliptic or lanceolate, acute, entire, 
some leaves at apex of sterile shoots with 2—5 teeth; petioles 0.5—2 mm 
long; flowering shoots 2—3 cm long; pedicels 6—12 mm long, to 1 7 mm in 
fruit; corolla 6— 7 mm in diameter; sepals triangular, short; follicles 
glabrous, the erect style arising from the dorsal side very near the apex. 
FL June, fr. July. (Plate XVII, Figure 1 1 ). 

Open slopes, rock crevices, rock streams. — E. Siberia: Dau. (E. part). 
Gen.distr.: Mong.: N. part of Mongolia adjacent to Dauria. Described 
from the Shilka River, below the mouth of the Daban River. Type in 
Leningrad. 

Note. In its habitus S. d ahur ic a Maxim, resembles S. a lp i na Pall., 
but is distinguished by broader leaves without glaucous bloom below, 
divergent fruiting sepals, and looser, broader inflorescences. 

Series 3. Alpinae Pojark. — Inflorescences terminating leafy shoots, 
corymbiform, dense; stamens longer than petals; fruiting sepals erect; 
style arising from dorsal side of follicle slightly below the apex. Leaves 
whitish below, smooth owing to papilliform epidermal excrescences. Buds 
with several pairs of free scales. Only 2 species in the Soviet Union. 

13. S. alpina Pall., Fl. Ross. I (1784) 35; Ldb., Fl. Ross. II, 13; Turcz., 
Fl.baic.-dah. I, 360; Kryl., Fl. Zap. Sib. VII, 1451 . - S. a lp in a var. 
altaica Maxim, in A. H. P. VI (1879) 183.- Ic: Pall., 1. c, tab. 20. 



229 



Shrub 50—80 cm high, devoid of pubescence, with slender, virgate, often 
arched -upcurved branches sometimes to 25—35 cm long, covered with 
strongly ramentaceous grayish brown bark; leaves dark green above, 
light and whitish below, lanceolate or lance -linear, short -acuminate, 
cuneately tapering toward base, without conspicuous petioles, 8—20 mm 
long, 1.5—4 mm broad, entire, on sterile shoots to 25mm long, 8 mm 
broad, sometimes denticulate; flowering shoots usually numerous, sometimes 
up to 25 on each branch, short, 1—3 cm, light brown; pedicels 2.5—6 mm 
long, to 10— 13 mm in fruit; corolla 5—6 (7) mm in diameter, with obovate 
„ qq or orbicular petals; flower buds not pigmented; follicles slightly hairy ventrally 
along the suture and at the apex, 2—3 mm long. Fl. June— July, fr. August. 
(Plate XVII, Figure 6). 

Mountains, forest zone, mainly in moist creek valleys, also in lower 
part of the high-mountain zone in alpine meadows and moss -and -lichen 
tundra; descends below the mountain-forest zone and grows on riverbanks, 
in forest clearings, rarely inbogs. — W.Siberia: Alt.; E.Siberia: Ang.-Say., 
Dau. Gen. distr.: Mong. — throughout the northern mountainous part 
of Mong., Tannu-Ola; Tibet and Central China. Described from Lake 
Baikal. Type in Leningrad. 

14. S.tianschanica A. Pojark. sp. nov. in Addenda VIII, p. 376. — 
S. oblongifolia Rgl. et Herd, in Bull. Soc. Nat. Mosc. XXXIX (1866) 42 
(non W. et K.).— S.crenifolia Rgl. in schedis. — S. alpina var. alt ai c a 
(non Maxim.) Fedtsch. in Consp. fl. turk. Ill (1909) 6. 

Low shrub, devoid of pubescence; branches 5—7 cm long, covered with 
peeling grayish-brown bark and with 1—5 axillary short flowering shoots 
1—2.5 cm long, with light brown or dark purple bark; leaves oblong obovate, 
6—20 mm broad, rarely narrower, obtuse or even rounded at the apex, 
mucronulate, cuneately tapering to very short petiole, glaucous -green 
above, whitish below, glabrous, rarely slightly hairy; pedicels 2.5—5 mm 
long; corolla 5— 6 mm in diameter; flower buds carmine, with persistent pink 
spots at flowering (always?); follicles glabrous. Fl.May— July, fr. August. 

Upper part of forest zone and lower part of high -mountain zone, on 
slopes, along streams and in alpine meadows. — Centr. Asia: T. Sh. (only 
E. part), Ketmen, Kurla, E.and-SE tip of Lake Issyk-kul'). Gen. distr.: 
Dzu.-Kash. — Kuldja. Described from the Terskei Ala-Tau. Type in 
Leningrad. 



Series 4. Trilobatae A. Pojark.— Flowers in umbellate inflorescences 
terminating leafy shoots; stamens shorter than petals; fruiting sepals erect; 
style arising from dorsal side of follicle very near the apex. Quite 
glabrous small shrubs, with broad leaves trilobate or coarsely dentate at 
the apex and distinctly but not prominently veined. 

15. S.trilobata L., Mant. II (1771) 244; Bge. in Ldb., Fl. Alt. II (l 830) 
214; Ldb., Fl. Ross. II, 1, 11; Fedtsch., Consp. fl. turk. Ill (l 909) 7 (excl. 
var.); Kryl., Fl. Zap. Sib. VII, 1487. - Ic. : Pall., Fl. Ross. I, 1 (1784) 
tab. XX. 

Shrub 15—100 cm high, devoid of pubescence, with broad, dense crown and 
grayish-brown branches; leaves with 2—5 mm petioles, orbicular or obovate, 



230 



cuneate or rounded at base, unequally dentate in upper part, usually 
shallowly trilobate, 6—20 mm long, 4—20 mm broad; flowering shoots 
yellowish or very light brown; pedicels (5)8— 18 mm long, with small 
linear bracts near the middle or lower; corolla 5— 8 mm in diameter; 
petals orbicular, slightly emarginate; follicles 2— 5 mm long, glabrous, very 
rarely slightly hairy along ventral suture and at the apex. Fl. June — July, fr. 
August. (Plate XVII, Figure 5). 

Open steppe slopes, mainly stony mountain slopes. — W. Siberia: Alt., 
Ang. -Say. (W. Sayans); Centr. Asia : Balkh. (E. part), Dzu. -Tarb - the SW 
border of the species passes through Tarbagatai. Gen. distr. : recorded 
for Jap.-Ch. — N.China (Chihli) and Korea. However, the Chinese specimens 
differ from the Altai ones, which casts doubt on the identity of the species. 
Described from Siberia. Type in London. 



Series 5. Pilosae A.Pojark.— Flowers in umbellate or corymbiform 
inflorescences terminating leafy shoots; stamens shorter than petals; 
fruiting sepals erect; styles arising from dorsal side near the apex. 
Pubescent shrubs; leaves oval-cuneate with very prominent veins. 
Several species growing in E. Asia (japan, China) and in the mountains 
of Middle Asia. 

16. S.pilosa Franch. in Ann. Sc. nat. ser. VI, XVI (1883) 282.- S. t r i - 
lobata var. pub e s c en s Rgl. in A. P. Fedchenko, Put. v Turk. No. 18 
(1882) 23; Fedtsch., Consp. fl. turk. Ill (1909) 7. 

Small, branching shrub 30— 70 cm high, with pubescent shoots; leaves 
4—20 mm long, 3—18 mm broad, oblong -cuneate or subrhomboid, unequally 
dentate above the middle, mostly trilobate, with very short (l— 3 mm) 
petioles, densely pubescent on both sides but more densely so below; 
flowering shoots 2—6 (l0)cm long, light brown, usually densely pubescent; 
inflorescences 6— 20 -flowered, umbellate or subcorymbiform; pedicels 
(3)6—16 mm long; corolla 7—10 mm in diameter, the petals erect, oboval; 
follicles about as long as sepals, inflated dorsally, with rounded apex, 
with sparse pubescence usually confined to the ventral suture and the 
apex. FL June — July, fr. August. (Plate XVII, Figure 4). 

Stony slopes, among rocks, in juniper or walnut forests, also on open 
slopes, at 1,000 — 2,500 m. — Centr. Asia: Pam.-AL — on N. slopes of the 
Alai Range (usually), one site is known from the district of Kafirnigan 
(Kanyaz); T. Sh. — continuous distribution area in W. Tien Shan (as far 
east as the Susamyr River) and one site in E. part of the Trans -Hi 
Ala-Tau (Karabulak). Endemic. Described from W. Tien Shan (Siemssen). 
Type in Paris. 



301 



17. S.pubescens Turcz. in Bull. Soc. Nat. Mosc. V (1832) 190; Kom. and 
Alis., Opred. rast. Dal'nevost. kraya II (1932) 627. — S.laucheana Koehn. 
in Mitt. Deutsch. Dendr. Gesellsch. VIII (1899) 56 (nomen nudum). 

Shrub 1 .6—2 mm high; shoots slightly ribbed, hairy, when young covered 
with very light brown bark; leaves obovateto oblong-rhomboid, dark green 
above, sometimes subglabrous when adult, grayish and rather densely hairy 
below, 7—2 mm long, 6—12 mm broad, on sterile shoots to 40 mm long and 



231 



302 



20 mm broad, with large, unequal teeth from the middle or only at the 
apex, sometimes nearly trilobate; flowering shoots 1— 3.5 cm long; 
inflorescences many -flowered, with glabrous pr slightly hairy pedicels; 
corolla 5—7 mm in diameter; follicles inflated dorsally, rounded at the apex, 
hairy only along the suture and at the apex. Fl. July, fr. September. 

Rocks and dry slopes.— Far East: Uss. — only along the Suifun River. 
Gen. distr.: Jap.-Ch. — Manchuria, N. China (Chihli, Shansi, and Kansu) 
and N. Korea. Described from the vicinity of Kalgan. Type in Leningrad. 



Series 6. Crenatae A. Pojark. — Inflorescences corymbiform, dense, 
terminating leafy branches; stamens longer than petals; fruiting sepals erect; 
style arising from dorsal side of follicle slightly below the apex. Leaves 
obovate, entire, more or less dentate only in certain species and only on 
sterile shoots. Apart from the USSR species there is also S.mongolica 
Maxim, in Mongolia and W. Tibet. 

18. S.crenata L., Sp.pl. (1753)489 (excl.syn.); Ldb., Fl. Ross. II, 1 1 ; 
M. B., Fl.taur.-cauc. I (1808) 392; Grossg., Fl. Kavk. IV (1934) 283. - 
S.hypericifolia a uralensis, e sawranica et I besseriana 
Ser. in DC. Prodr. II (1825) 543.- S. sawranica Bess, ex DC, I.e. 
(pro synon). — S.hypericifolia a latifolia et /3 longifolia Ldb., 
Fl. Alt. II (1830) 214.- S.ob longifolia Ldb., Fl. Ross. II, 13 (non W. et K.). 
et K.). - S. cr enifolia C. A. M. in Beitr. Pflzk. Russ. Reich. VI (1844) 
43; Shmal'g., Fl. I, 315; Fedtsch., Consp. fl. turk. Ill (l 909) 6 ; Poyarkova 
in Fl. Yugo-Vost. V (1931) 4 86.— S.crenifolia var. capitata Trautv., 
A.H. P. II (1873) 528.-S.crenifolia a pallasiana et j3 capitata 
Maxim, in A. H. P., VI (1879) 180.- S. cr enifolia f. glaberr ima Korsh., 
Mem. Acad. Petersb. VII Ser., VII (1898) 149.- S.crenifolia var. p alias ii 
f. puberula Litw. in Sched. HFR (1901), No. 873.- S.crenifolia var. 
pallasii f.glabrata Litw., 1. c, No. 874. — Ic. : Ldb., Ic. pi. Fl. ross. V 
(1834) tab. 428,429; Fedch. and Fler., Fl. Evrop. Ross. (1910), Fig. 415; 
Fl. Yugo-Vost. V, Fig. 418.- Exs.: HFR, No. 873, 873a, 874. 

Shrub ca. 1 m high, with finely ribbed, pubescent shoots; leaves usually 
puberulent, rarely glabrous, on sterile shoots 1.5— 3 (3. 5) cm long, 0.3— 1 .5 mm 
broad, oblong-obovate to cuneate or obovate, entire or denticulate or crenate 
from the middle, less often nearly from base, sometimes with only the apex 
dentate, usually with 3 prominent longitudinal veins; leaves on flowering 
shoots smaller, 0.6— 2 cm long, and narrower, oblong-obovate or lanceolate, 
entire, with 1 midrib and 1 or 2 pairs of lateral veins; petioles 1—5 mm 
long; flowering branchlets 1— 8 cm long; inflorescences 10— 12 -flowered, 
compact, sometimes hemispherical (f. capitata Maxim. ), with pubescent 
pedicels only 1.5—2 times as long as flowers; follicles 2— 3 mm long, 
pubescent, less often subglabrous, x / 4 - Vs longer than the oblong sepals. 
Fl. May, fr. July. (Plate XVIII, Figure 2). 

Forest -steppe zone, thickets of steppe shrubs, forest edges, steppe 
sands, fields, boundaries, high areas in river floodplains, less often on 
open stony and silty slopes, rarely on limestone slopes. — European part: 
V.-Kama, M. Dnp., V.-Don, Transv., Bl., L.Don, L. V.; W. Siberia: Ob - only 
in SW part and at one site on the Irtysh River in the former Tara County, 



232 



U. Tob., Irt., Alt.; Centr.Asia: Ar. -Casp. (N. part),Balkh.; Caucasus: 
Cisc, Dag., E. and S. Transc. Gen.distr.: E.Europe. Described from 
W. Siberia. Type in London. 

Economic importance. Used for fastening of sands; ornamental and 
nectariferous. 

19. S. lasiocarpa Kar. et Kir. in Bull. Soc. Nat. Mosc. XV (1842) 536; 
Fedtsch., Consp. fl. turk. Ill (l 909) 6. — S. hy p e r i c i f o 1 i a (non L.) Kar. et 
Kir., 1. c, p. 348. — S. c r e nat a var. s u b 1 ob at a Rgl. et Herd., Enum. pi. 
Semen, in Bull. Soc. Nat. Mosc. XXXIX (1866) 86. - S. c r e n i f o 1 i a var. 
integrifolia Rgl. in sched. 

Shrub; shoots glabrous or yellowish-tomentose, slightly angled, with light 
brown, later peeling bark; branches long, sometimes to 30—40 cm, with 
numerous crowded flowering axillary shoots, the upper 1—2 cm long, the 
others ca. 3 cm long, the lowermost sometimes 4—4.5 cm long; leaves 
oblong, oboval, or elliptic, cuneately tapering to 1.5—3 mm petiole, glabrous 
or slightly hairy on the margin (usually in lower part); leaves on sterile 
shoots 10—22 (28)mm long, 2.5—6 mm broad, sometimes with 3 teeth 
at the apex; leaves on flowering shoots entire, 5—12 (15) mm long, 2—6 mm 
broad; inflorescences dense, the lower sometimes with ramified 
2— 6 -flowered branchlets; pedicels 2— 5 mm long, 4— 8 mm long in fruit; 
flowers 7—10 mm in diameter, with rounded or rounded-oval petals 2—4 mm 
long; follicles 3—4 mm long. Fl. May— July, fr. from July. (Plate XVIII, 
Figure 1 ). 

Mountain forests and lower part of the high -mountain zone, alpine 
meadows. — Centr. Asia: Dzu. -Tarb. (S. part), T. Sh. — in E. part in the 
Trans -Hi and Kirghiz Ala-Tau (formerly Aleksandrovskii) ranges, 
Pam.-Al. — N. slope of the Alai Range. Endemic. Described from the 
Dzungarian Ala-Tau (Sarkan). Type in Leningrad. 

20. S. ferganensis A. Pojark. sp. n. in Addenda VIII, p. 377. — S. lasio- 
carpa Franch. in Ann. d. sc. nat. VI Ser.,XVI (1883) 283 (non Kar. et Kir.). 

Shrub; shoots slightly angled, light brown, glabrous or with a dense 
yellowish tomentose pubescence; leaves on sterile shoots 20— 37 mm long, 
5 — 12 mm broad, oblong, elliptic, acute, or else oboval and obtuse or rounded 
at the apex, glabrous or slightly hairy on the margin in lower part, sometimes 
with 2 or 3 teeth at the apex; leaves 6—11 on flowering shoots, (10) 12—22 mm 
long, 3— 7 mm broad; flowering shoots few, 4— 12 per branch, 5— 8 cm long, 
only the uppermost shorter, 3— 4 cm; inflorescences loose, 6— 12 -flowered; 
pedicels of upper flowers 4—6 mm long, others 8— 18 mm long; flowers 
large, 10—12 mm in diameter, with rounded -oval petals 4—5 mm long and 
broad; follicles 4—4.5 mm long. Fl. July, fr. from August. 

Undergrowth of broadleaf mountain forests. — Centr. Asia: T. Sh. : 
Fergana Range, Andizhan and Namangan mountains. Endemic. Described 
from the Aflatun River in W. Tien Shan. Type in Leningrad. 

Series 7. Hypericifoliae A. Pojark.— Flowers in sessile umbels; fruiting 
sepals erect; style arising from dorsal side of follicle very near the apex. 
Leaves on sterile shoots differ from those on flowering shoots in the 



233 



presence of teeth at the apex, sometimes also in shape. Buds ovate, 
with free scales.— Apart from the 2 species occurring in the Soviet Union, 
the W. European S. o b o v a t a W. et K. and S.hupehensis Rehd. from 
Central China also belong to this series. 

21. S.hypericifolia L., Sp.pl. (1753)489; Ldb., Fl. Ross. II, 12 ; M. B..F1. 
taur.-cauc. I (1808) 392; ShmaPg., Fl. I, 315; Poyarkova in Fl. Yugo-Vost. 
V (1931) 487; Kryl., Fl. Zap. Sib. VII (l 933) 1451 . - S. a mb i g ua Pall., 
Fl. Ross. I (1784) 36.- S. cr enata (non L.) Pall., Verz. Pflanz. Taur. (1796) 
108; C. A. M., Enum.pl. cauc.-casp. (1831) 166.- S.acutifolia Willd., 
Enum. pi. hort. Berol. I (1809) 540.— S.hypericifolia |3 plukenetiana 
et 7 ac uta Ser. in DC, Prodr. II (1825) 543.- S. hyper icifolia 7 
brevi folia Ldb., Fl. Alt. II (l 830 ) 215.- S.hypericifolia a genuina 
et microphylla Ldb., Fl. Ross. II (1844) 12.— S. hyper icifolia var. 
acutifolia Dipp.,Handb. d. Laubh. Ill (1893) 464.- S.hypericifolia a 
typica Maxim, in A. H. P. VI (187 9) 178 (excl. syn.).- S.hypericifolia 
var.heterophy 11a Somm. et Lev. in A. H. P. XVI (1900) 143.— 
S. hypericifolia var. turkestanica Ldb. ex Beissn., Schelle et Zab., 
Handb. d. Laubholzb. (1903) 149.— S. hypericifolia var. obovata et 
304 morpha subalpina Zinserl. in Acta fl. ross. I (1915) 19.— S. obovata 
Grossh., Fl.cauc. IV (1934) 283 (non W. et K.).- Ic. : Ldb., Ic. Fl. Ross. V 
(1834) tab. 430; Vol'f and Pal., Opr. der. i kust. (l 904), figure on p. 373.- 
Exs. : HFR, No. 872, 872a. 

Shrub 50—150 cm high; branches light brown, often long, virgate, with 
numerous, crowded sessile umbels; young shoots glabrous or tomentose- 
pubescent; leaves 10— 25 mm long, 1.5 — 8 mm broad, glabrous or puberulent 
when young, oboval, oblong -elliptic or lanceolate, obtuse or acute, entire, 
those on sterile shoots sometimes with 2—5 teeth at the apex, cuneately 
tapering to short, 1.5— 5 mm petiole; inflorescences 4— 10 -flowered; 
pedicels glabrous or slightly pubescent, 5—15 mm long, to 18 mm long in 
fruit; flowers 5—8 (9) mm in diameter, with oboval or ovate petals and 
triangular sepals 1 / 3 — l j 2 as long as follicles, the latter glabrous or pubescent. 
Fl. May —June, fr. from July. 

Steppe and forest -steppe zone where, together with other steppe shrubs, 
it forms thickets; also gully slopes and open, often also stony slopes; 
in mountain regions of Central Asia in the shrub zone, on open slopes, in 
juniper woods, and on mountain riverbanks; in the Caucasus in shrubthickets 
on mountain slopes, penetrating to alpine meadows where it grows as a 
small, low, much branched shrub, often with broader leaves (morpha 
subalpina Zinserl.)- — European part: V. -Kama, V. -Don, Transv., BL, 
Crim., L. Don, L. V.; Caucasus: Cisc, Dag., W., E., and S. Transc; 
W. Siberia: U. Tob., Irt., Alt.; E. Siberia: Ang. -Say. — only the westernmost 
part; Centr.Asia: Ar. -Casp., Balkh., Dzu. -Tarb., Mtn. Turkm., T. Sh., 
Pam.-Al. Gen. distr.: Arm. -Kurd.— Turkish Armenia, Dzu. -Kash. — 
Kuldja, and Mong. — NW Mong. Described after a cultivated specimen. 
Type in London. 

Note 1. Grossheim cites S. ob o vat a W. et K. for the Caucasus. This 
species, distinguished from S.hypericifolia by larger flowers, longer 
pedicels, and absence of stomas on upper surface of leaves, grows wild only 
in France and Spain; in the Caucasus it is replaced by S.hypericifolia L., 
which varies more in the Caucasus and the mountain regions of Central 



234 



306 



Asia than it does in the European part, and which often produces larger 
flowered specimens than the typical plants, which were apparently taken 
for S. o b o v a t a W. et K. 

S. hy p er i c if ol ia X S.crenata. — Hybrids between these 
species occur rather frequently in the Caucasus (Dag., E. and 
S.Transc.) and rarely (L. Don, V. -Don) in the European part. In 
their morphological characters they approach one or other of these 
species. They can easily be recognized by the structure of their 
inflorescences, which are often sessile umbels on one branch in the upper 
part, lower down corymbs on a more or less developed leafy shoot, as in 
S.crenata but with longer pedicels; leaves on sterile shoots resemble 
those of S. hy p e r i c if o 1 ia but with lateral veins approximate at 
base and more or less parallel to the midrib as in S.crenata, with 
dentation also similar to that of S. c r e n a t a. 

22. S. aquilegifolia Pall., Reise III (1776) Anh. 734.- S.thalictroides 
Pall., Fl. Ross. I (1784) 34; Turcz., Fl. baic. -dah. I, 360.- S. hy p e r i c i f o 1 i a 
var.thalictroides Ldb., Fl. Ross. II, 1 (1844) 13. — S. hy p er ic if ol j. a 
(non L.) Turcz., 1. c, 359. - Ic. : Pall., Reise, III, tab. P., f. 3; Ej., Fl. Ross. I, 
tab. 18. 

Rather low shrub with slender, erect, light brown or grayish -brown 
branches; leaves glabrous or minutely velutinous -pubescent, more densely 
so below, at base of inflorescence 2—10 mm long, 1.5—4.5 mm broad, 
oboval, with rounded apex and cuneate base, entire or with 3 teeth at the apex, 
to 17 mm long and broad on sterile shoots, only the lowest oboval, the others 
broadly cuneate, with few, mostly 3—5 large lobelike teeth at the apex; 
inflorescences 2—5 (7)-flowered; pedicels glabrous, 2—5 (6) mm long; 
flowers 6—8 mm in diameter; follicles glabrous or slightly pubescent, covered in 
lower V 4 — Y 3 by erect sepals. Fl. June, fr. July. (Plate XVIII, Figure 3). 

Open dry steppe slopes.— E.Siberia: Dau. (except E.part). 
Gen. distr.: Mong. — mountain districts, NE and Centr. Mong. and N. China 
(Muni-ula and Alashan ranges). Described from Dauria. Type in London. 

S. aquilegifolia X S. alp in a. — The hybrid of these species from 
the vicinity of Verkhneudinsk [Ulan-Ude] does not differ from S. alpina 
in the flower structure; the inflorescences are as in S. alpina except 
that the uppermost are subsessile; leaves glaucous below (owing to 
papilliform epidermal excrescences as in S. a lp in a), oboval; leaves on 
sterile shoots with large teeth. Another hybrid from Dauria closely 
resembles the first in its leaves, but the inflorescences are more similar 
to those of S. aquilegifolia. 

S. aquilegifolia X S. media. — The hybrids approach one or other 
of the species; inflorescences corymbiform on very short shoots, sometimes 
partly sessile as in S. aquilegifolia but with long pedicels; flowers 
mostly large, as in S. media; leaves intermediate in size and shape, but 
in type of dentation resembling S. m e d i a. Rarely in Dauria. 



Genus 719. SIBIRAEA MAXIM. 

Maxim. in A. H. P. VI (1879) 213. 

Dioecious shrubs with simple, entire exstipulate leaves; inflorescence 
paniculate. Staminate flowers with 20—25 stamens and very small, little 

* From the word Siberia. 

235 



307 




PLATE XVIII Is ■ 



>-■»"..«.. °» ulga , ^ a 'r, e , r " e *""• W 416 4-s.bir.,.' 



al ta 



6- A.k 



a L., leaf 



on sterile shoot; 



lensisOaxm.) C.K. Schn 
' amtschaticus Rydb. 



5773 



236 



developed pistils arranged at base of hypanthium; pistillate flowers with 
5 (4—7) pistils connate at base along ventral suture, with short stamens, 
the anthers not containing pollen; ovules 4—6, anatropous; fruit an 
aggregate follicle; follicles glabrous, erect, mostly with 2 large seeds. 

Apart from the USSR species, there are S. croatica Degen in Croatia 
and Herzegovina and S.tomentosa Diels.in central China. 

1. Inflorescence axis and hypanthium glabrous; follicles 5— 5.5 mm 
long, 2—2.5 mm broad; leaves large, to 1 1 cm long, 2.2 cm broad, 
obtuse 1. S. altaiensis (Laxm.) C. K. Schn. 

+ Inflorescence axis and hypanthium hairy; follicles 3.5—4 mm long, 

1—1 .5 (2) mm broad; leaves small, to 6 cm long, 7 mm broad 

2. S. tianshanica (Krassn.) A. Pojark. 

1. S. altaiensis (Laxm.) C. K. Schn., 111. Handb. Laubholz. I (1906) 486. - 
Spiraea altaiensis Laxm. in Nov. Coram. XV (1771, juni) 554. — 
Spiraea laevigata L., Mant. II (1771, octob.) 224; Ldb., Fl. Alt. II (1930) 
216; Fl. Ross. II, 1, 15.- Spiraea altaica Pall., Reise II (1773) App. 739; 
Fl. Ross. I (1784)37.-Sibiraea laevigata Maxim, in A. H. P. VI (1879) 
215; Kryl., Fl. Zap. Sib. VII, 1458.- Ic: Laxm., 1. c, tab. 29, f. 2; Pall., I.e., 
(1784) tab. 23. 

Erect, smooth shrub 60—150 cm high; branches thick, covered with dark 
brown bark; leaves entire, thickish, gray -green, lanceolate, 30— 40 mm 
long, 7—20 mm broad, rounded at the apex, finely mucronulate, cuneately 
tapering downward, glabrous or ciliate -margined when young; inflorescences 
paniculate, developing at ends of long lateral shoots of the current year and 
composed of 5—10 simple racemes 3—10 cm long arising from leaf axils, 
diminishing upward; flowers white; inflorescence axis and pedicels glabrous: 
staminate flowers 6— 7 mm in diameter, the pistillate slightly smaller, 
with pedicels 2—6 mm long, with linear or lanceolate bracts at base; the 
hypanthium broadly campanulate, smooth outside; follicles glabrous, 
5—5.5 mm long, 2—2.5 mm broad, greatly exceeding the erect sepals. 
Fl. June, fr. from July. (Plate XVIII, Figure 4). 

Open mountain valleys, mountain slopes; sometimes forming large 
thickets. W.Siberia: Alt. Endemic. Described from the Altai. Type 
in London. 

Economic importance. Cultivated as an ornamental plant. The leaves 
are used by local populations as a substitute for tea. 

2. S. tianschanica (Krassn.) A. Pojark. sp. n. in Addenda VIII, p. 237. — 
Spiraea laevigata var. t i an s c hani c a Krassn., Sp. rast. vost. Tyan'- 
Shanya (1887)49.- S. alt ai ens i s var. 1 a nc if o 1 i a M. Pop. in sched. 

Low shrub to 0.5 m high, with thick branches; leaves narrowly lanceolate, 
sessile, acute, with a short cartilaginous cusp, gradually tapering downward, 
20—60 mm long, 4—7 mm broad, glabrous, sometimes slightly ciliate -margined; 
panicle of 4—6 spiciform racemes, 2.5—7 cm long; flowers tightly crowded 
on thick axis, the lower with pedicels 0.5—3 mm long, the others sessile, 
the staminate 5 mm in diameter; pistillate flowers unknown; inflorescence 
axes, pedicels, and hypanthium rather densely pubescent; follicles glabrous, 
3.5—4 mm long, 1—1. 5 (2) mm broad. Fl. June, fr. August. 



237 



310 



Meadows in the subalpine zone.— Centr.Asia: T. Sh. Described from 
the Kokzhar River. Endemic. Type in Leningrad. 



Genus 720. ARUNCUS ADANS. 

Adans.Fam.pl.il (1763) 295. 

Dioecious perennials with thick, lignifying rhizomes and compound bi- 
or tripinnate exstipulate leaves; flowers small, in spiciform racemes 
arranged in panicles; sepals 5, adnate at base to the flat hypanthium; 
petals 5; stamens 15— 30, with filaments several times as long as petals; 
stamens of pistillate flowers with short filaments and imperfectly developed 
anthers; pistils 3—5, alternating with sepals, as small rudiments in staminate 
flowers; follicles cartilaginous, dehiscing by ventral suture; seeds 
bacilliform, with endosperm. 

1. Flowers in spiciform racemes up to 3 cm long, these in turn arranged 
the shape of a loose, simple raceme; leaves ca. 10 cm long, 

biternate 4. A. parvulus Kom. 

+ Inflorescence paniculate, composed of numerous longer racemes; 

leaves bipinnate, with 3—9 leaflets 2. 

2. Staminate flowers 4—5 mm in diameter, twice as large as the pistillate; 
petals oblong (2.5—3 times as long as broad); inflorescences mostly 
crowded, few -branched 3. A. kamtschaticus Rydb. 

+ Staminate flowers 3—4 mm in diameter, slightly larger than the 
pistillate; petals oboval (twice as long as broad); inflorescences 
usually luxuriant, profusely branching 3. 

3. Racemes of pistillate flowers dense, spiciform 

2. A. asiaticus A. Pojark. 

+ Racemes of pistillate flowers sparse 1. A. vulgaris Raf. 

1. A. vulgaris Raf., Sylva Tell. (1838) 152.- Spiraea ar uncus L., Sp. 
pi. (1753) 490; Ldb., Fl. Ross. II, 1, 16 (ex parte). - U 1 m a r i a aruncus 
Hill., Hort. Kew. (1769) 214. — Aruncus Sylvester Kostel. in Ind. hort. 
Prag. (1844)5 (nom. nud.); Shmal'g., Fl. I, 316; Grossg., Fl. Kavk. IV (l 934) 
283.-Astilbe aruncus Trev., Bot. Zeit. XIII (l 855) 814. - A. s i 1 v e s t e r 
a vulgaris (ex parte) Maxim, in A. H. P. VI (1879) 1 70. — Sp ir a e a 
paniculata St. -Lag., Ann. Sc. bot. Lyon, VII (1880) 135.— A. aruncus 
Karst., Deutsch. Fl. (1882) 779; Rchb., Ic. Fl. Germ. XXIV (1909) 146.- 
Ic: Hegi, 111. Fl. Mitteleur. IV, 2, tab. 145, f. 16 et f. 1004. 

Perennial, 1—2 m high, with thick, woody rhizome developing numerous 
shoots; leaves long-petioled, to 1 mm long, bipinnate, usually with 9 leaflets, 
the upper leaflets entire, short -petioled, the lower with conspicuous petioles 
to 5 cm long, pinnatipartite into 3—7 lobes, the lower lobes usually tripinnate; 
lobes oblong or lanceolate -oval, the terminal broader, tapering to a long sharp 
point, cuneate or truncate, rarely cordate, mostly oblique at base, glabrous 
or sparsely hairy along the veins, biserrate; panicles compound, very 
large, spreading, to 50 cm long, individual racemes to 15 cm long; racemes of 
staminate flowers dense, spiciform, those of pistillate flowers sparse; 

* Latin name of this European species, which was called Barb a cap re a (goatsbeard) by pre-Linnaean 
authors. 



238 



staminate flowers 3— 3. 5mm in diameter, with oval or slightly spatulate 
petals, 1— 1.5mm long, 1 mm broad; pistillate flowers 2.5— 3 mm in diameter; 
follicles 2.5 —3 mm long, 1 mm broad, glabrous. Fl. from mid -June to 
beginning of August, fr. from mid-August. (Plate XVIII, Figure 5). 

Forests up to the subalpine zone. — Caucasus : Cisc, Dag., W., E., and S. 
Transc. Gen. distr.: Centr. Eur., as far east as Volhynia and Lithuania. 
Described from Austria. Type in London. 

Economic importance. Ornamental plant cultivated in Europe since the 
17th century. Leaves and flowers were formerly used in medicine as an 
antipyretic. Cyanogenetic glycosides are also found in leaves and flowers. 

2. A. asiaticus A. Pojark. sp. n. in Addenda VIII, p. 377. — Spiraea 
aruncus Ldb., Fl. Ross. II, 1, 16 (ex parte); Turcz., Fl. baic. -dah. I, 312. — 
A. Silvester a vulgaris Maxim, in A. H. P. VI (1879) 170 (ex parte).— 
A. Silvester Kom., Fl. Mansh. II (1904) 461; Kom. and Alis., Opred. rast. 
Dal'nevost.kraya II (1932) 628.- Ic. : Kom. and Alis., 1. c, p. 188, Fig. 1 . - 
Exs. : Karo, PL Amur, et Zeaen., No. 335. 

Perennial, 1—2 mm high, with glabrous or slightly hairy shoots; leaves 
bipinnate, resembling the preceding species in size and type of dissection; 
leaflet lobes broadly ovate to lanceolate -oval, glabrous or with scattered 
hairs mainly along the veins below, truncate, broadly cuneate, less often 
cordate at base, usually abruptly narrowed to a sharp point, biserrate; 
panicles to 35 cm long; racemes of staminate and pistillate flowers dense, 
spiciform, to 10 cm long; staminate flowers 3— 4 mm in diameter, the 
pistillate slightly smaller, 2.5—3 mm; petals oboval, ca. 1.5 mm long, 1 mm 
broad; follicles 2.5—3 mm long, 1.5 mm broad, glabrous. Fl. June— July, 
fr. from August. 

Deciduous and sparse mixed forests and forest edges.— E.Siberia: 
Lena-KoL, SE part and Dau.; Far East: Okh. (S. part), Ze.-Bu., Uda, Uss., 
Sakh. Gen. distr. : Jap.-Ch. (Manchuria, N. Korea). Described from Ol'ga 
outpost, on the coast of the Sea of Japan. Type in Leningrad. 

Economic importance. Ornamental plants. 

3. A. kamtschaticus Rydb. in N. Amer. Fl. XXII, 3 (1908)256.- 
Spiraea aruncus Ldb., Fl. Ross. II, 1, 16 (ex parte). — Aruncus 
Silvester a kamtschatica Maxim, in A. H. P. VI (1879) 170.— 
Aruncus Silvester (non Kostel.) Kom., Fl. Kamtsch. II (1929) 234; 
Hulten, Fl. kamtch. Ill (1929) 43. 

Perennial, 30—150 cm high, with glabrous or slightly hairy shoots; 
leaves bipinnate, with 5—7 leaflets, the upper leaflets entire, the lower 
pinnate, 3 - or 5-lobed, the lower lobes sometimes tripinnate; lobes 
broadly lanceolate to lanceolate -oval, glabrous, cuneate or truncate, usually 
oblique at base, tapering to a more or less long tip, biserrate or doubly 
incised, serrate; panicles 6—15 cm long, mostly compact, slightly branching, 
composed of tightly crowded racemes 2—8 cm long, rarely more luxuriant 
(to 2 5 cm long); in the low -growing alpine variety (f. alpihus Kom.); 
inflorescences much impoverished, sometimes reduced to a simple 
spiciform raceme; staminate flowers 4— 5 mm in diameter, with oblong - 
obovalpetals 1 .5—2 mm long, 0.5— 0.75 mm broad; pistillate flowers ca. 2.5 mm 
in diameter; follicles 2—3 mm long, 1—1.5 mm broad. Fl. from July to 
mid -August, fr. September. (Plate XVIII, Figure 6). 



239 



312 



313 



InBetula ermani forests, alder groves, dry meadows, and grassy 
slopes. — Arctic: Arc. Sib. (known only at the Lena River mouth), An.; 
Far East: Kamch. , Okh. (N. part). Gen.distr.: Jap.-Ch. — Kurile Islands; 
Ber. — Aleutian Islands, Alaska. Described from Kamchatka (from the 
vicinity of Petropavlovsk). Type in New York. 

4. A.parvulus Kom. in Bull. Jard. Bot. Ac. Sc. URSS, XXX (1932) 203; 
Kom. and Alis., Opred. rast. Dal'nevost. kraya (1932) 62 8. 

Perennial; rhizome woody, simple; stems 5—20, short, erect, simple, 
15—30 cm high, with 2—5 basal leaves, 5 — 8 cm long, the petioles as long 
as or somewhat longer than the blade, mostly bi- or tripinnate or else 
bipinnate with lower leaflets tripinnate and the upper leaflets 5-lobed; 
lobes oval -rounded, 1—4 cm long and broad, glabrous or slightly hairy, 
with very prominent veins below; inflorescence sometimes composed 
of a single dense, spiciform raceme, but usually of several short racemes 
to 3 cm long collected into a sparse, simple raceme; staminate flowers 
ca. 3 mm in diameter, with lanceolate bracts; pistillate flowers unknown; 
follicles 2—3 mm long, ca. 1 mm broad. Fl. from end of May, fr. August. 

Limestones, rock crevices.— Far East: Uss., along the Suchan River. 
Endemic. Described from Mt. Zolotoi in the Suchan River valley. Type 
in Leningrad. 



Genus 721. SORBARIA A.BR. 

A.Br.in Aschers., Fl. v. Brandenb.( 1864) 177. 

Flowers in terminal, mostly large panicles, bisexual, with 40—50 stamens 
and 5 (4—8) pistils opposite to sepals; ovary connate along the ventral 
sutures to the middle or slightly beyond, with several ovules; seeds few. 
Shrubs with pinnate leaves and persistent stipules. 

Approximately 10 species distributed throughout Asia. 

1. Fruit drooping; follicles 2— 3 mm long 1. S. olgae Zinserl. 

+ Fruit erect; follicles at least 5 mm long 2. 

2. Flowers 7—11 mm in diameter, in pyramidal panicles; stamens twice 

as long as petals 2. S. sorbifolia (L.) A. Br. 

+ Flowers 12—16 mm in diameter in obovoid panicles; stamens as long 

as or slightly longer than petals 3. 

3. Ferruginous bristly -glandular pubescence characteristic; leaflets to 

7 cm long, 4 cm broad, in closely approximate pairs 

3. S. rhoifolia Kom. 

+ Glandular pubescence absent; leaflets to 4 cm long, 2 (2.5) cm broad, 

their pairs freely arranged 4. S. pallasii G.Don. 

Series 1. Lindleyanae A. Pojark.— Fruits nodding; follicles small, 
2—3 mm long; panicles pyramidal, large, with pubescence of only simple 
hairs. 

Middle Asian species: S. lindleyana Maxim. (Himalayas), S. g i 1 g i - 
tens is Zinserl. (Kashmir), S. a r b o r e a C. K. Schn. (Central China) and 
S.angustifolia Zbl. (Afghanistan). 

* Referring to the resemblance to the leaves of Sorbus aucuparia L. 



240 



1. S. olgae Zinserl. in Not. Syst. Herb. Hort. Petr. VI, 2 (1926) 33.- 
S.sorbifolia var. glabra Maxim, in A. H. P. VI (1879) 223 (ex parte). - 
Basilima sorbifolia Fedtsch., Consp. fl. turk. Ill (1909) 8 (non Rafin.). 

Shrub with glabrous light shoots; leaves 10— 20 cm long, with linear, 
acute stipules; leaflets 7—9 pairs, oval-lanceolate, abruptly narrowing to 
a sharp point, 2.3—5 cm long, 0.8—2.3 cm broad, glabrous above, with single 
hairs along the veins below, biserrate; panicle 15—25 cm long, many- 
flowered, dense, with glabrous axis and pedicels; flowers unknown; follicles 
2—3 mm long. 

Gorges. — Centr. Asia: Pam. -AL, on N. slope of the Alai Range near 
Shakhimardan. Described from there. Endemic. Type in Leningrad. 



Series 2. Sorbifoliae A.Pojark.— Fruits erect; follicles 5 mm long; 
flowers in pyramidal panicles; stamens twice as long as petals; pubescence 
of simple branching and short glandular hairs. Apart from the USSR 
species there belong here closely related but distinguishable forms from 
central China. 

2. S. sorbifolia (L.) A. Br. in Aschers., Fl. v. Brandenb. (1864) 177; 
Maxim, in A. H. P. VI (1879) 22 (f. glabra (ex parte) et st ellipila); 
Kryl., Fl. Zap. Sib. VII, 1459 (f. glabra et s t e 11 ip i 1 a); C.K.Schn., 
Handb. Laubh. I (1906) 488 (a.typica). -Spiraea sorbifolia L., 
Sp.pl. (1753) 490; Se'r. in DC, Prodr. II (1825) 545; Turcz., Fl. baic. -dah. I, 
36; Ldb., Fl.Ross. II, 1, 15; S.pinnata Moench, Meth. (1794) 663. - 
Basilima sorbifolia Raf., New. Fl. Amer. Ill (l 836) 75. — S c h i z o n ot u s 
sorbifolius Lindl. in Steud., Nomencl. II, 2 (1841) 531.— Sorbaria 
stellipila C. K. Schn., Handb. Laubh. I (1906) 489 (var. typ i c a et 
insert a); Kom., Fl. Kamch. II (1929) 233; Kom. and Alis., Opred. rast. 
Dal'nevost. kraya II (1932) 62 9. — S o r b ar i a assurgens hort. ex 
C. K. Schn., 1. c, 490. - Ic. : Pall., Fl. Ross. I, tab. 24. 

Tall shrub, 1—3 m high, with abundant coppice roots and puberulous, 
rarely glabrous shoots; leaves long-elliptic, 12— 25 cm long, 6— 13cm 
broad, with petioles 2—5 cm long, the 9—21, mostly 17 lanceolate acuminate 
leaflets 2.5—8 cm long, 0.8—2.5 cm broad, glabrous, less often with simple 
or branching yellow hairs along the veins below, biserrate; stipules 
broadly cordate to linear, dentate or entire; panicles 12—30 cm long, 
5—12 cm in diameter; inflorescence axis, pedicels, and often lower part 
of hypanthium usually outwardly puberulous with admixture of short glandular 
pubescence or else more or less densely covered with bushy rufous hairs 
and then the leaves usually similarly pubescent along the veins below; 
flowers 7—11 mm in diameter, with suborbicular petals; stamens twice as 
long as petals; petals densely hairy; follicles pubescent, 5 mm long. 
Fl. from mid -June to beginning of September, fr. from August. 

Banks of mountain and forest streams and small rivers; locally 
forming large, very dense thickets; rather sparse coniferous, mixed, and 
deciduous forests, forest edges, bog margins.— W. Siberia: Ob — E. part, 
between 59 and 56.6°N.,as far west as 82.5° E. E.Siberia: Lena-Kol. (S. half) 
(S. half), Ang. -Say. (N. part), Dau.; Far East: Kamch., Okh., Ze. ~Bu., Uda, 



241 



Uss., Sakh. Gen. distr.: Mong. — N. Mongolia; Jap. -Ch. —Manchuria, 
Korea, Japan. Apparently, the Chinese plants should be separated as a 
species. Described from Siberia. Type in London. 

Economic importance. Ornamental shrub, firmly established in 
cultivation, one of the most widespread shrubs in the parks and gardens 
of the Soviet Union, escaping in some places. 

Note. S.stellipila C. K. Schn. is recorded for the vicinity of 
Vladivostok. Schneider indicates this species for Japan, as opposed 
to the continental S. sorbifolia (L.) A. Br., having densely pubescent 
ovaries, pedicels, and lower part of hypanthium. Schneider distinguishes 
between S . s t e 1 1 i pi 1 a var. ty p i c a — with leaves densely pubescent 
below with bushy hairs — and var. insert a with glabrous leaves. 
Examination of abundant herbarium material has failed to confirm the 
presence of differences between the Japanese and the continental plants as 
indicated by Schneider: both plants are characterized by the densely 
pubescent ovaries, pubescent pedicels, and often also pubescent hypanthium; 
specimens with only slightly pubescent ovaries occur very rarely. 
Pubescence of bushy hairs can also not be considered as a specific 
character, since specimens with such pubescence are distributed — though 
irregularly — throughout the distribution area of S. sorbifolia: in the 
western part of the area they are rare, whereas in the eastern part (in 
S. Yakutia and the Far East) they are quite common. Therefore we do not 
find sufficient grounds for recognizing S. stellipila C. K. Schn. as a 
separate species, and we regard it as a variety of S. sorbifolia (L.) 
A.Br. 



Series 3. Pallasianae A. Pojark. —Fruits erect; follicles 6 — 8 mm long; 
„ 1 (. panicles ovoid; stamens as long as or slightly longer than petals. Pubescence 
of simple, bushy, and glandular -bristly hairs. Two species in the Soviet 
Union. 

3. S. pallasii (G. Don) A. Pojark. comb. n. — Sp i r a e a sorbifolia var. 
alpina Pall., Fl. Ross. I (1784) 38; Ldb., Fl. Ross. II, 16. - Sp i r a e a 
grandiflora Sweet, Hort. brit. (182 7) 194 (nom. nud.). — Sp. pallasii 
G. Don, Gard. Syst. II (1832) 520; Rgl. et Til., Flor. ajan. (1858) 80. - 
Sorbus grandiflora Maxim. inA.H. P. VI(1879)223; Kom ., Fl. Mansh. II 
(1904)469. -Sor bus al p in a Dipp. Laubh. Ill (1893) 503; Kom. and Alis., 
Opred. rast. Dal'nevost. kraya II (1832) 528. — Basilima alpina Zbl. 
Strauch. Spir. (1893) 115.- Ic: Pall., Fl. Ross., I.e., tab. 25; Kom. and Alis., 
1. c, tab. 189. 

Low shrub, to 40 cm high, the divaricate branches covered with peeling 
reddish-gray bark, with smooth brownish shoots usually — like the petioles 
and inflorescence axes — puberulous, rarely glabrous, or, if densely covered 
with bushy yellow hairs, then inflorescence axes, pedicels, and mostly 
underside of leaves with similar pubescence; leaves 4—15 cm long, 
their 9—13 (l5)leaflets linear -lanceolate to oval-lanceolate, 1—4 cm 
long, 0.2—1.2 (2) cm broad, glabrous or sparsely hairy or with dense 
pubescence of bushy hairs; inflorescence 2—8 cm long, 2—6 cm in 
diameter; bracts 5— 8 mm long, the lower broadly oval, the upper narrowly 
lanceolate; corolla 12—15 mm in diameter, the hypanthium pubescent 



242 



outwardly; stamens slightly exceeding petals; follicles 6— 7 mm long, 
hairy. Fl. July, fr. September. (Plate XIX, Figure 7). 

Balds, cliffs, and stony taluses in the alpine zone.— E. Siberia: 
Lena-Kol. (S. part); Far East: Okh. (S. part), Ze. -Bu. (N. part), Uda, 
Uss. (N. part). Endemic. Described from the northern shore of Lake 
Baikal. 

4. S. rhoifolia Kom. in Bull. Jard. Bot. XVI (1916) 174; Kom. and Alis., 
Opred. rast. Dal'nevost. kraya (l 932) 628. — Ic. : Kom. and Alis., 1. c, tab. 
190. 

Small, densely leafy shrub; thick shoots covered — like petioles and 
inflorescence axis — with a pubescence of two kinds — of short simple 
hairs and of long, rufous, glandular -bristly hairs; leaves to 15 cm long, 
with 9—11 (13) closely arranged sessile leaflets 2.5— 7 cm long, oval - 
lanceolate, gradually acuminate, with oblique base, glabrous above, densely 
whitish-pubescent with bushy hairs — mainly when young — below, biserrate; 
panicles dense, rounded -ovoid, to 8 cm long, 6 cm in diameter; flowers 
large, to 15 mm in diameter, pinkish white, with suborbicular petals, with 
outwardly glandular hypanthium and sepals; follicles 8 mm long, appressed- 
hairy. Fl. June, fr. August. 

Dry stony slopes.— Far East: Uss. — on riverbanks (Botchi and 
Svetlaya rivers). Endemic. Described from the Svetlaya River. Type 
in Leningrad. 



Genus 722. SPIRAEANTHUS * MAXIM. 
Maxim, in A. H. P. VI (1879) 226. 

Shrub with narrowly linear pinnate exstipulate leaves, the bisexual 
flowers 5 (4)-merous; hypanthium broadly campanulate; petals firm, 
late -deciduous; stamens 20—25; pistils 2—5, connate at base along ventral 
suture, opposite to sepals; ovules 2, anatropous; seeds mostly 1. Only 
1 species known. 

1. S. schrenkianus (Fisch. et Mey.) Maxim, in A. H. P. VI (1879)227; 
Fedtsch., Consp. fl..turk. Ill (1909) 8. — Spiraea schrenkiana Fisch. 
et Mey.,Ind. sem. hort. Petrop. IX (1842) 96 (nomen nud.). 

Low shrub with broad, dense crown; shoots puberulous, covered with 
grayish -yellow bark splitting longitudinally and peeling; leaves 2 — 13 cm 
long, 0.2—0.5 cm broad, narrowly linear, pinnate, with thickened petioles 
and 20—35 pairs of thickish oval leaflets 1—2 mm long, covered like the 
petiole with a fine pubescence; inflorescences in the shape of oblong sparse 
panicles 9—20 cm long, developing at ends of new annotinous shoots and 
composed of one simple and more axillary racemes, up to 1.5 — 10 cm 
long, diminishing upward; flowers pink, aromatic, with lanceolate bract 
and bracteole, the pedicelsl .5— 2mm long; hypanthium broadly campanulate, 
covered outwardly with sparse pubescence and yellowish scaly glands; 
fruiting sepals erect, obtuse, 2 / 5 as long as follicles; follicles gray -hairy, 
5 mm long. Fl. June, fr. August. (Plate XIX, Figure 6). 

* From the Greek sp i r a e a, meadowsweet and a nth os, flower. 



243 



Slopes of gorges (in the Kara-Tau Mountains) and on banks of takyrs* 
in the Golodnaya Step), on gravelly soils; usually forming thickets. — 
Centr. Asia: Balkh. (SE part), T. Sh. - only in the Kara-Tau Range, 
Syr D. (N. part). Endemic. Described from the Chu River in the Golodnaya 
Step. Type in Leningrad. 

Economic importance. Ornamental shrub; the fine, dense, pinkish-brown 
wood may be used for making small articles. 



Genus 723. EXOCHORDA * LINDL. 

Lindl.in Gard. Chron.f 1858) 925. 

Shrubs with leaves entire or dentate near the apex, exstipulate; flowers 
„ _ in racemes, large, white, 5 -merous, polygamo-dioecious or monoecious; 
hypanthium obconical; sepals caducous in fruit; stamens 25 (in USSR 
species) or 15 ; pistils 5 (much reduced in staminate flowers); ovary with 2 
almost completely connate ovules; fruit capsulelike, consisting of 5 laterally 
flattened woody follicles dehiscing by suture and each containing one winged seed , 

1. Fruits 12-1 7 mm long 1. E. alberti Rgl. 

-t- Fruits 8— 10 (12) mm long 2. E. tianschanica Gontsch. 

1. E. alberti Rgl. in A. H. P. VIII (1884) 6 96.- E.korolkovii Lav., 
Arb. Segrez. (1885) 39; Fedtsch., Consp. fl. turk. Ill (190 9) 9.- Albert ia 
simplicifolia Rgl., Ind. Sem. hort. Petr. 1883 (nom. nud.). — E.grandi- 
flora var. alberti Aschers. et Graebn., Syn. mitteleur. Fl. VI (1900) 30. — 
Ic: Rgl., 1. c, tab. XIII. 

Tall shrub to 4 m high, profusely branching, with glabrous, slender, red- 
brown young shoots and grayish-brown branches; leaves light green, 
glabrous, oblong -oboval or lanceolate, cuneately tapering toward base, 
the lower sessile, the others petiolate, 2—10 mm long (leaves on sterile 
shoots sometimes oblong -ovate), short -acuminate or obtuse, with short 
cartilaginous mucro; racemes 3— 10 -flowered, 3— 8 cm long, glabrous, the 
flowers crowded along inflorescence axis; hypanthium with 10 prominent 
ribbed nerves; sepals short, rounded -triangular; corolla 2.8— 4.5 cm in 
diameter; stamens 25, approximate in 5's, opposite to sepals; styles free, 
erect or reflexed; fruits 12— 17 mm long, the pedicels 1— 3 mm long, those 
of lower fruits sometimes to 5—10 cm; seeds flat, oval with winged margin. 
Fl. April — June, fr. from August. (Plate XIX, Figure 5). 

Undergrowth of sparse, broadleaf forests, slopes among shrubs. — 
Centr. Asia: Pam.-Al. — E. part. (Gissar, Bal'dzhuan, Karategin, Kulyab, 
Darvaz; only exceptionally penetrating further west than Karatag). 
Endemic. Described from the Yakhsu River. Type in Leningrad. 

Economic importance. Ornamental shrubs. 

2. E. tianschanica Gontsch. in Sched. herb. Fl. As. med., fasc. XXI -XXIII 
(in Acta Univ. As. Med. ser. VIII, Botanica, fasc. 17) (1934) 91.- Exs.: Herb. 
Fl. As. Med., No. 574. 

* [Clay-soil sections in desert and semidesert areas.] 

" From the Greek exos, exterior and chorda, cord, referring to the fact that the receptacle forms a free 
subulate excrescence protruding between the follicles. 



244 



Very closely related to the preding species, from which it is distinguished 
by fruit, 8—10 mm in length and diameter, sometimes only to 12 mm; in 
addition, flowers and fruit mostly more remote on axis, and the axis more 
slender in fruit; flowers more numerous in racemes, up to 13. Fl. May and 
beginning of June, fr. from August. 

Undergrowth of sparse broadleaf forests, slopes among shrubs.— 
Centr.Asia: T. Sh. : Chatkal and Fergana ranges. Endemic. Described 
from the Yassy River in the Fergana Range. Type in Leningrad. 

Economic importance. Ornamental shrubs. 



Subfamily 2. POMOIDEAE Focke in Engl. u.Pr., Nat.Pfl. Ill, 3 (1888) 18.- 
Pomaceae Lois .-Deslongch. Manuel d. PI. Usuelcas, I (1819) 211 . — 
Hypanthium cyathiform, ascidiform or nearly tubular. Sepals and petals 5; 
stamens mostly 20, less often 5 or 10, sometimes 25 or more. Carpels 
2 — 5, united with inner wall of the hollow inflorescence axis and mostly 
also connate; inflorescence axis becoming fleshy toward the apex, forming 
a pseudocarp. — Trees and shrubs with simple, lobate or imparipinnate 
leaves; stipules more or less conspicuous, mostly deciduous. Flowers 
in umbellate, racemose or paniculate inflorescences. 

1. Pseudocarp with nutlets (endocarp stony) 2. 

+ Pseudocarp without nutlets (endocarp coriaceous) 5. 

2. Flowers solitary, ca. 5 cm in diameter 732. Mespilus L. 

+ Flowers in more or less many -flowered inflorescences, small. ... 3. 

3. Leaves entire; branches unarmed 724. Cotoneaster Medik. 

+ Leaves dentate or lobate, at least at the apex; branches mostly with 

spines . 4. 

4. Leaves small, crenate -serrate 731. Pyracantha Roem. 

+ Leaves larger, lobed 733. Crataegus L. 

5. Flowers solitary, terminal; leaves entire, short -petioled 

725. Cydonia Mill. 

+ Flowers in more or less many-flowered inflorescences; leaves dentate, 
lobate or pinnatipartite, less often entire but then petioles more or 
less long 6. 

6. Leaves evergreen; inflorescences densely rufous -tomentose, in the 
shape of a compact pyramidal panicle *Eriobotrya Lindl. 

+ Leaves deciduous; inflorescence not as above 7. 

7. Flowers in simple racemes; petals oblong, cuneately tapering 
downward „ 731. Amelanchier Medik. 

+ Flowers in umbellate or corymbiform inflorescences; petals rounded 
or oval, more or less short -clawed 8. 

8. Leaves entire or slightly lobate, with 8—12 pairs of parallel, prominent 
veins terminating in teeth; calyx caducous in fruit (fruits quite smooth 
at the apex) 72 9. Micromeles Decne. 

+ Leaves pinnately compound, if entire or lobate then with veins 
disappearing in mesophyll near the margin; fruits usually with 
persistent calyx 9. 

9. Leaves pinnate or lobate; flowers 1.2—2 cm in diameter; shoots 
unarmed 728. Sorbus L. 



245 



320 



+ Leaves entire or, if deeply partite, then shoots very spiny; flowers 

(2)2.5— 4.5cm in diameter 10 

10. Styles free at base; fruit pulp with grit cells 726. Pyrus L 

+ Styles connate at base; fruit pulp without grit cells . . .727. Malus Mill 



G e nu s 724. COTONE ASTER * M ED IK . * * 
Medik.,Gesch.d.Bot.(1793) 85. 

Flowers small, in racemose or corymbiform inflorescences, rarely 
fascicles of 1—3; stamens 20; ovary composed of 2—4 (5) carpels adnate to 
the hypanthium by the dorsal side. Pomes small, mealy, with 2—4 nutlets 
sunken to 2 / 5 — 2 /s in the pulp and more or less covered above by persistent 
sepals. Unarmed shrubs, very rarely small trees with entire, alternate 
leaves. 

Economic importance. Most species are ornamental shrubs valued for 
their beautiful shape and the bright color of their fruit; they also have 
the advantage of developing well on scarps and stony slopes, even on 
open, sunny slopes. 

1. Flowers open, with white spreading petals; styles (nutlets) 2; fruits 
erect. (Section C hae nop e t a lum) 2. 

+ Flowers not open, crown-shaped, with erect, more or less pinkish 
petals; styles (nutlets) mostly 3, sometimes 2 or 4. (Section Ortho - 
petalum) 7. 

2. Flowers 2—4 in leaf axils or in a short raceme; ovary, hypanthium, 

and sepals sparsely appressed -pubescent . . . 5. A. oligantha A. Pojark. 
+ Flowers usually in compound, rarely in simple corymbs 3. 

3. Inflorescence and lower surface of leaves glabrous or rather sparsely 
pubescent (not tomentose!) 6. C.multiflora Bge. 

+ Inflorescence with at least axes and pedicels tomentose; leaves 

tomentose below 4. 

4. Ovary, hypanthium, and sepals sparsely appressed -pubescent; leaves 
oblong, oval, or elliptic 9. C.taurica A. Pojark. 

+ Ovary, hypanthium, and sepals densely white -tomentose; leaves usually 
large 5. 

5. Fruits black 6. 

+ Fruits red; leaves 0.5—2 cm long and broad, rarely larger 

8. C. racemiflora (Desf.) C.Koch. 

6. Fruits large, 7— 9 mm long; inflorescence 10— 20 -flowered, to 3.5 cm 

in diameter; leaves on short shoots 2—5 cm long, 1.5—4.8 cm broad . . . 

7. C. insignis A. Pojark. 

+ Fruits smaller, 6— 8 mm in diameter; inflorescence 6— 12 -flowered; 

leaves on short shoots 0.9— 2.5 cm long, 0.4— 1 .8 cm broad 

10. C. saxatilis A. Pojark. 

7. Flowers in racemes or corymbiform panicles; fruits black 8. 

+ Flowers 1—3 in axillary fascicles 9. 

8. Leaves tomentose below; fruits with glaucous bloom 

l.C. melanocarpa Lodd. 

* From the Latin cotonea, cotoneum.on Malum cotoneum — the Roman name of quince 

(Cydonia), apparently owing to the resemblance between the quince leaves and the cotoneaster leaves. 
** Treatment by A.I. Poyarkova. 



246 



+ Adult leaves glabrous below or sparsely appressed -pubescent; fruits 
without bloom, lustrous 2. C.lucida Schlecht. 

9.^ Leaves glabrous below or sparsely pubescent; flowers solitary, 

rarely in pairs 4. C.uniflora Bge. 

+ Leaves white -tomentose below; flowers in groups of 1—3(4) 

3. Cintegerrima Medik. 



Section 1. ORTHOPETALUM Koehne, Dendr. (1893) 224.- Petals erect 
at anthesis, usually pink. 



Series 1. Melanocarpae A.Pojark. — Fruits black, covered by waxy 
bloom; leaves tomentose below. 

1. C.melanocarpa Lodd., Bot.Cab.XVI (1829) sub tab. 1531; Voronov in 
Fl. Yugo-Vost.; V (1931) 488.- Mespilus cotoneaster L., Sp.pl. 
(1753) 479 (ex parte); M. B., Fl. taur. -cauc. I, 388; Wahlenberg, Fl. Gothland 
Syn. (1837) 17.- C. vulgaris (non Lindl.) Ldb., Fl. Ross. II, 1, 92 
(ex parte). — M e sp i lu s cotoneaster var. nigra Ehrh., Beitr. IV (1789) 
19; Wahlberg, Fl. Gothoburg. (1820)53.— C. integerrima var. f r uc t o 
nigro Medik., Gesch. d. Bot. (1793) 85. — C. vulgar i s J3 melanocarpa 
Bge. in Ldb., Fl. Alt. II (1830) 219; Litw. in Sched. herb. Fl. Ross. V (1905) 
98. — C. vu lgar i s var. h a e m at o c a r p a Rupr., Fl. ingr. (l 853) 350; Litw., 
1. c. — C. nigra Fries, Sum. Veg. Scand. I (1846) 175.— C.polonica 
Jastrz, ex Rostaf., Fl. Polon. prodr. (1872) 121.- C. nigra Rgl. in A. H. P. 
11(1873)315 (excl.var.); Zinserl. in Izv. Gl. Bot. Sada XXIII (1924) 13.- 
C. integerrima var. melanocarpa Kryl.,Fl. Zap. Sib. VII (1933) 1461. — 
Ic: Pall. Fl. Ross. I (1784) tab. XIV (right-hand figure).- Exs.: HFR, 
Nos. 1469 and 1470. 

Shrub to 2 m high; shoots more or less tomentose when young; 
annotinous shoots glabrous, lustrous, red-brown; petioles 1—3 (5) mm long; 
leaves ovate or elliptic, emarginate, rarely acute, dark green and slightly 
hairy, rarely glabrous above, whitish-tomentose below; flowers in groups 
of (3)5—15, rarely numerous in nodding axillary racemes or corymbiform 
panicles, the axis hairy, sometimes tomentose, the pedicels less pubescent, 
sometimes subglabrous; hypanthium glabrous or only slightly pubescent; 
stamens 20, with linear filaments, abruptly narrowed at the apex; styles 
mostly 3, less often 2; fruits obovoid -globose, 7—9 mm long, brownish red 
when immature, at maturity black with glaucous bloom; nutlets 2 or 3, the 
style arising in upper part. Fl. June, fr. September. 

Mountain regions of the USSR; mainly on stony slopes, open or overgrown 
with shrubs and forests, in the middle mountain zone, rising locally to the 
subalpine zone; in the plains of the distribution area — in open forests and 
groves, also mainly on stony soils.— European part: Kar.-Lap., Dv.-Pech., 
Lad.-Ilm., U. V., V. -Kama, U.Dnp., M. Dnp., V.-Don., Transv.; Caucasus: 
all districts except Talysh; W. Siberia: Ob (S. part), U. Tob., Irt., Alt.; 



247 



322 



E.Siberia: Lena -Kol. (S. part), Ang. -Say., Dau.; Centr.Asia: Ar.-Casp. 
(N. part), Balkh., Dzu.-Tarb., T. Sh. (only in the Talass Ala-Tau and the 
Fergana Range), Pam. -Al. (only on N. slopes, of the Alai Range). Gen. distr.: 
Centr. Eur., Dzu.-Kash. (Kuldja), N.Mong., Jap. -Ch. (Manchuria, rarely). 
Described from Dnepropetrovsk (formerly Ekaterinoslav). Type unknown. 

Economic importance. Ornamental shrub widespread in gardens and 
parks of the Soviet Union; various hybrids of C. m e 1 an o c a r p a with other 
Cotoneaster species may also be of interest for cultivation. 

Note. C.melanocarpa varies greatly in size, shape of leaves and 
their pubescence (especially above), size and pubescence of axial parts, 
size of fruits and density of their glaucous bloom. However, the examination 
of abundant herbarium material did not permit determination of the 
relationship between these characters and the distribution area. 

This species is sometimes cited for the Soviet Union under the erroneous 
name C. nigra Wahlenberg (or Wahlberg); these 2 combinations should, 
however, be excluded even from the synonyms of C.melanocarpa, for 
they were reported by the above-mentioned author due to a misunderstanding. 

C.melanocarpa naturally hybridizes with a series of other 
Cotoneaster species, and some of these hybrids are quite common. 

C.melanocarpa X C.multiflora Bge. — Hybrids often occurring 
in the Caucasus and very often in Central Asia (Dzungarian Ala-Tau, E. 
and Centr. Tien Shan). These hybrids are very frequent and varied as to the 
character of combinations of parent species, either occupying an intermediate 
position between them or closer to one than the other. A number of them 
are described under binomial specific names. The following belong here: — 
Cigna va E. Wolf (izv. Lesn. Inst. XV (1907) 240) (Central Asia), which 
is fully intermediate between the parent species, having dichasially 
branching inflorescences with pink corolla more open than in C.melano - 
c ar pa, with brown-red, nearly black fruit, and the leaves pubescent 
below. — C.pseudomultiflora M. Pop. (E-yull. Mosk. Obshch. Est., nov. 
ser. XLIV (1935) 127) from the vicinity of Alma-Ata, with densely pubescent 
lower surface of leaves, many-flowered compound corymbiform 
inflorescence, white open corolla, and globose dingy purple -red fruit.— 
C.submultiflora M. Pop. (l. c. 126) (vicinity of Alma-Ata) is more 
closely related to C.multiflora than the preceding: leaves slightly 
pubescent below, fruits globose and bright red. The following apparently 
also belong to the group of hybrids of C.melanocarpa X multiflora: — 
C.megalocarpa M. Pop. (i.e. 128) (Alma-Ata), with large leaves 
pubescent below, abundant fruiting, and large vinaceous-red fruits 10—11 cm 
in diameter; — C. po ly ant h e m a E. Wolf (Mitteil. d. deutsch. dendr. 
Gesellsch. 1924, p. 325), from the Alma-Ata vicinity, very closely related 
to C.melanocarpa, from which it is distinguished by its many- 
(11 — 25-flowered) inflorescence, its white, divaricate petals, smaller 
leaves of flowering branches, very irregular ripening of fruits, which 
are lustrous black with slight glaucous bloom, and a vegetative period one 
month longer than that of C.melanocarpa. The indications of earlier 
authors also pertain to these hybrids: C.laxiflora (non Jacq.) Bong, et 
Meyer in Suppl. II, Fl. alt. (1841 ) 190, and also C. vulgaris ]3 erythro- 
carpa Ldb. Fl. Alt. II (1830) 219; Kryl., Fl. Alt. 426.- C . integer rim a var. 
erythrocarpa et var. gl ab r at a Kryl., Fl. Zap. Sib. VII, 1461 et 
C.multiflora |3 pubescens Rgl. in A. P. H. II (1873) 316. 



248 



323 



C.talgarica M. Pop. (l. c. 126) (W. Talgar River, near the Bogdanovich 
Glacier) very closely related to C.melanocarpa, but distinguished from 
it by narrower leaves 2 / 3 as long and less tomentose below, by the 
few-flowered, slightly pubescent inflorescence, and by its black fruits lacking 
glaucous bloom; apparently a hybrid of C. m e 1 a no c ar p a Lodd.X C.uni- 
f lora Bge. (p. 25l). 

C.melanocarpa Lodd. X C.integerrima Medik. (p. 2 50). 

C.melanocarpa Lodd. X C. r a c e mi f lo r a C. Koch, occurring in 
the Caucasus (C. nigr a. var. daghestanica Zinserl. in Izv. Gl. Bot. 
Sada XXIII (1924) 15) as well as in Central Asia, usually with tomentose 
inflorescences either resembling in structure those of C. m e 1 an o c a r p a 
or else corymbiform as inC.racemiflora, erect or more or less nodding; 
fruits with usually 2, less often 3 nutlets. 



Series 2. Lucidae A. Pojark. — Fruits black without waxy bloom, with 
1—4 (2) nutlets; leaves slightly hairy below. 

2. C.lucida Schlecht.in Linnaea (1854) 541; C. K. Schneid., Illustr. 
Handb.d. Laubh. I (1906) 750.- C.acutifolia Lindl. in Ldb., Fl. Ross. II, 

1 (1844) 92 (non Turcz.).— C. nigra var. a c ut i f o 1 i a Wenz. in Linnaea 
(1884) 183.- Ic: C. K. Schn., 1. c, f.421 c,423c- Exs.: HFR, No. 360. 

Erect shrub to 2 m high; shoots densely appressed -pubescent when 
young; leaves 1.7—5 (6) cm long, 0.8—3.5 (4) cm broad, with hairy petioles 
2—6 mm long, the blade dark green and lustrous, glabrous or with hairs 
along the midrib above, yellowish-tomentose below only when young, later 
light, yellowish, slightly appressed-pubescent; flowers 5—12 in loose 
corymbiform racemes shorter than leaves; inflorescence axes and 
pedicels with more or less dense yellowish appressed pubescence; ovary 
and hypanthium glabrous or slightly hairy; sepals broadly triangular, 
villous -tomentose on the margin, %— % as long as the pink petals; stamens 
20, the filaments gradually narrowing; styles 3 (4); fruits black, lustrous, 
without bloom, ovoid -globose, 7—9 mm in diameter, with usually 3, rarely 

2 nutlets, rather densely hairy at the apex and bearing style in upper part. 
Fl. June, fr. September. (Plate XIX, Figure 2). 

Rocky slopes, shrub thickets and sparse larch -and -mixed forests, river 
pebbles.— E. Siberia: Ang.-Say.— only at southern end of Lake Baikal. 
Endemic. Described after a cultivated specimen. Type in Berlin. 

Economic importance. Very ornamental plants owing to lustrous 
foliage and abundant fruiting. Tolerant of clipping, therefore used for 
borders and hedgerows. 



Series 3. Integerrimae A. Pojark.— Fruits red, with 3 or 4 nutlets; 
petals as long as or scarcely longer than sepals. 

3. C.integerrima Medik., Gesch. d. Bot. (1793) 85 (ex parte); 
Kihlm. in Meddel. Soc. pro fauna et fl. fenn.XIV (1900) 114; Grossh., 
FL cauc. IV (1934) 284.- Mespilus cotoneaster L., Sp, pi. (1753) 
479 (ex parte); M. B., Fl. taur. -cauc. I, 388. - C. vu lg a r is Lindl. in 
Trans. Linn. Soc. XIII (1821) 101; Ldb., Fl. Ross. II, 1, 92 (ex parte); 
Zinserl. in Bull. Jard. bot. d. Republ. Russe, XXIII (1924) 12. - Ic: Pall. 
Fl.Ross. I, 1 (1784) tab. XIV (middle and left-hand figures). 

249 



Erect shrub with wide spreading crown, not more than 1 .5 m high; shoots 
densely appressed -pubescent when young; leaves on flowering shoots 
0.9—4 cm long, 0.4—2.7 cm broad, broadly ovate, less often oblong -ovate, 
short -acuminate, with cartilaginous cusp, light green at the apex, dull, less 
often sparsely hairy above, whitish or grayish-tomentose below; flowers 
mostly in pairs, less often solitary or 3(4) in nodding raceme, always 
_„. shorter than leaves, with slightly villous -tomentose axis and pedicels and 
with glabrous hypanthium; stamens 20, styles 3 or 4; fruit purple -red 
at maturity, from subglobose to ovoid or obovoid, 8—11 mm long, with 
(2) 3 or4nutlets, hairy at the apex; style arising above the middle, 
less often from the middle. Fl. June, fr. from September. 

Stony mountain slopes, taluses and rocks.— European part: Crim.; 
Caucasus: Cisc.,W., S. (and rarely E.) Transc. Gen. distr.: Eur. (to the 
Baltic States, S. Scandinavia, and S. Finland). Described from Europe. 

Economic importance. Not widely cultivated in the Soviet Union; most 
indications (usually as C. vulgaris) should be referred to C.melano - 
c ar p a. 

Not e. Occurs in the Soviet Union only in the Crimea and the Caucasus; 
all reports concerning the European part, Siberia, and Central Asia are 
erroneous and refer partly to C.uniflora Bge., partly to C.racemiflora 
(Desf.) C. Koch, but mostly to C.melanocarpa Lodd. which, when its 
fruits are immature, is often not differentiated from C. integerrima 
or else is reported as one of its forms (C. vulgaris v.haematocarpa 
Rupr.), which is none other than C. me lanoc arp a Lodd. These two species, 
however, may be clearly differentiated in the various phases of growth: 
C.melanocarpa has a different inflorescence structure, with well 
developed axis and long pedicels, more open flowers, larger and brighter 
petals; the brown-red color which characterizes the immature fruits of 
C.melanocarpa has not been observed in C. integerrima fruits 
at any stage of maturity, and the tomentose pubescence on the lower 
surface of leaves is looser and consists of longer hairs. 

C. integerrima Medik. X C. m e la no c ar p a Lodd. —Hybrids 
occasionally occur in the Caucasus (Cisc, Dag., Armenia). In shape 
and leaf pubescence these hybrids approach either C. integerrima or 
C.melanocarpa; inflorescences with well developed axis as in 
C.melanocarpa, but few- (3— 5)-flowered, with more or less developed 
tomentose pubescence; fruits dark red or black, without glaucous bloom, 
small, nodding, with usually 3, less often 2 or 4 nutlets; style arising 
above the middle. 

C. integerrima Medik. X C.racemiflora C. Koch. — Hybrids are 
apparently rare (vicinity of Shemakha and Sardar-Bulak); leaves small, 
close to one or the other species in shape and pubescence of leaves; 
fruiting poor; fruits in twos or threes, axillary or in corymbs, with 
tomentose pedicels, erect or nodding (even on the same branch), red, 
with 2 nutlets, narrower than in C.racemiflora, the style arising in 
upper part. 

4. C.uniflora Bge. in Ldb., Fl. Alt. II (1830) 220; Ldb., Fl. Ross. II, 1, 
191; Kihlman in Meddel. Soc. pro fauna et flora fenn.XIV (1900) 114; 
Kryl., Fl. Zap. Sib. VII, 1462.— C.integerrima var. unifiora 



250 



325 



C. K. Schneid., Illustr. Handb. Laubh. I (1906) 747. - Ic. : Ldb., Ic. pi. Fl. Ross. 
Ill (1831) tab. 269; Kihlman, 1. c, f. 5, 6, 9.- Exs.: PI. Finland, exs., No. 752. 

Low shrub, 30—40 cm high, spreading, branching, sometimes quite 
prostrate and appressed to the ground; young slender shoots covered with 
yellowish appressed hairs; leaves with short, 2.5 mm petioles, the blade 
oblong-ovate, broadly ovate, or elliptic, acute or subobtuse, sometimes 
emarginate, often with a cartilaginous cusp, dark green and glabrous above, 
yellowish, lighter, glabrous or rather sparsely hairy below, 1—3 (4) cm 
long, 0.6—2.5 cm broad; flowers in axils of leaves solitary, less often 
in pairs, with nodding, glabrous pedicels shorter than or as long as 
flowers; hypanthium glabrous; sepals broadly ligulate, obtuse, ciliate- 
margined; petals greenish white or pinkish, scarcely longer than sepals; 
stamens 20; styles 3 or 4; fruits ovoid -globose, 6— 8(10) mm in diameter, 
purple -red or orange -red, with (2) 3 or 4 nutlets; nutlets subglabrous at 
the apex, the style arising mostly from below the middle, sometimes from 
the middle. Fl. June, fr. August — September. (Plate XIX, Figure 1 ). 

Stony slopes, rock streams, rocks, river pebbles, less often sedge -and - 
lichen tundra and alpine meadows, mostly in the alpine, less often in the 
forest zone. In the Altai it descends to open stony slopes.— Arctic; 
Arc. Eur.; European part: Kar.-Lap. (apparently only Kola Peninsula), 
Dv. -Pech., V. -Kama (Northern and Central Urals); W.Siberia: Alt.; 
Centr.Asia: Dzu. -Tarb., T. Sh. - E.,Centr., and W. (Talass Ala-Tau), 
and Pam.-Al. (single sites in Zeravshan, Karategin, Darvaz, Shugnan, 
and on N. slopes of the Alai Range). Gen. distr. : Chinese Dzungaria, 
Mong. — Tuva ASSR, NW Mongolia. Described from the Altai. Type in 
Leningrad. 

Note. It is difficult to ascertain on the sole basis of herbarium 
material— specimens from a few districts being quite insufficient — 
whether there is only one species or whether several very closely related 
geographic races should be separated. 

It should be noted that the monocarpous Cotoneaster from the 
Kola Peninsula and the Urals has, on the average, broader and larger 
leaves and larger fruit than the Altai specimens; it is unclear whether 
there are differences in color of petals; as a rule these are purple -red 
in Siberian plants, whereas orange -red petals predominate in northern 
plants (only the latter are known in the Khibiny mountain region) and 
yellow -fruited specimens (var.lutea Fries) occur. In the Pamir -Alai, 
in addition to subglabrous specimens indistinguishable from those from 
Altai, there are specimens with dense, almost tomentose pubescence 
below, which has been observed neither in the Altai nor in the Kola 
Peninsula. 

C.uniflora Bge. X C.melanocarpa Lodd. = C. nigra X pauci- 
f lor a Rgl. in A. H. P. II (1873) 315.).- Hybrids of these species are 
not infrequent in the Altai and the Sayans, where they vary considerably, 
showing various combinations of the characters of the two species. 
They usually differ from C.melanocarpa in more impoverished, 
often 1—3 -flowered raceme, but with more developed axis and pedicels 
than in C.uniflora. The pubescence on the underside of the leaves 
varies from slight to tomentose, and the leaf shape is closer to one or 
other of the species. The Kola Peninsula and Ural hybrids have more 



251 



(327) 




PLATE XIX. 1-Cot 



252 



329 



constant characters: in habitus they closely resemble C.uniflora, 
with raceme 1— 2 (3)-flowered, but the axis and pedicels are longer; 
leaves resemble those of C. uniflora but rather densely pubescent below; 
fruits dark red, with slight glaucous bloom. In Kar.-Lap.the forms of 
hybrid origin (apparently hereditary) penetrate much further south 
than C.uniflora ,(to northern tip of Lake Onega). In the literature they 
have been reported as C. n igr a (Kihlman, 1. c.) and C.integerrima 
var. nigra (Fl. Finland, exs., No. 753). 



Series 4. Oliganthae A. Pojark. —Fruits red with 2 (l) nutlets; petals 
1.5—2 times as long as sepals. 

5. C. oligantha A. Pojark. in Notul. syst. Inst. Bot. Ac. Sc. URSS, VIII, 
7-9 (1938). 

Shrub with young, sometimes also annotinous shoots covered with 
greenish-gray tomentum of rigid appressed hairs; bark of older shoots 
dark brown; leaves bright green above, with sparse appressed hairs, 
greenish-gray-tomentose below, mostly rounded at the apex, less often 
acute, sometimes emarginate, often mucronate, 8—17 mm long, 4—12 mm 
broad, on sterile shoots to 27 mm long, 19 mm broad; petioles tomentose- 
pubescent, 2—4 mm long; flowers on very short lateral branchlets in leaf 
axils, almost in fascicles of 2—4 or in short erect racemes half as long 
as leaves; axis very short, 2— 3 mm long, tomentose-pubescent; pedicels 
2—5 mm long, also tomentose; ovary, hypanthium, and sepals scattered - 
hairy to subglabrous; sepals broadly triangular, obtuse or subacute, with 
tomentose -fimbriate, mostly purple margin; corolla 8 mm in diameter 
with somewhat divaricate petals; stamens 20; styles 2; fruits with erect 
pedicels, red, subglobose, 8 mm in diameter, with 2 nutlets 4—5 mm long, 
3— 4 mm broad, flat ventrally, convex dorsally, with short pelta and with style 
arising almost at the apex. Fl. May— June, fr. August — September. 

Stony mountain slopes.— Centr.Asia: Balkh., Dzu. -Tarb., T. Sh. : Centr. 
(Kirghiz Ala-Tau) and W. (Talass Ala-Tau and Chimgan). Endemic. 
Described from the Arkatskie Mountains in Kazakhstan. Type in Leningrad. 



Section 2. CHAENOPETALUM Koehne, Dendr. (1893) 224. - Petals 
spreading at anthesis, white. 



Series 1. Multiflorae A. Pojark. — Fruits red; inflorescence profusely 
branching; all plant parts glabrous or slightly pubescent. 

6. C.multiflora Bge. in Ldb., Fl. Alt. II (1830) 220; Ldb., Fl. Ross. II, 1, 
93; M. Popov, in Tr. Prikl. Bot., Gen. i Sel. XXII, in 3 (1929) 446; Kryl., Fl. 
Zap. Sib. VII, 1463; Grossg., Fl. Kavk. IV (1934) 284. - C. r ef 1 ex a Carr., 
Revue hort. (1871) 520.- Ic: Ldb., Ic.pl. Fl. Ross. Ill (1931) tab. 254. 

Erect shrub 0.5— 1.5 m high; shoots with greenish yellow fine tomentose - 
pubescence when young, later glabrous, lustrous, reddish brown; leaves with 
rather long (5—10 mm) petioles, obovate or elliptic, mostly broad, usually 



253 



330 



obtuse, emarginate, less often — especially on sterile shoots — short - 
acuminate, rounded or broadly cuneate at base, dark green and glabrous 
above, glabrous below from the very beginning or with initially rather 
dense, fine appressed pubescence later becoming sparse, 1.5—4.5 (5)cm 
long, 1.2—3.5 (4)cm broad; flowers in rather loose, erect, 6— 20 -flowered, 
compound, dichasially branching corymbs, glabrous or with slightly 
pubescent axis, pedicels, and hypanthium; axes and pedicels slender; 
sepals broadly triangular, ciliate at the margin and often reddish; corolla 
ca. 1 cm in diameter, with orbicular, spreading petals; styles 2; 
inflorescence erect in fruit; fruits bright red, oblong -obovoid to subglobose, 
6—10 mm long, 3—7 mm broad, with 2 (rarely 1 ) nutlets 3—5 mm long, 
2.5—4 mm broad; style arising slightly below the apex. Fl. June, fr. from 
August. 

Rocks and stony mountain slopes, among shrubs.— Caucasus: Cisc, Dag., 
S.and W. Transc; W.Siberia: Irt. (S..), W. Alt.; Centr.Asia: Dzu.-Tarb., 
T. Sh., Mtn. Turkm. — Kopet-Dagh (rarely), Pam.-Al. (rarely — single sites 
in Zeravshan, Shugnan, and N. slopes of the Alai Range). Gen. distr.: 
central China: Kansu Province. Described from the Chingiz-Tau Mountains 
in Kazakhstan. Type in Leningrad. 

Economic importance. Rarely cultivated in the Soviet Union, but 
deserving attention, being more ornamental than C.melanocarpa 
which is cultivated in the Soviet Union; fruits more brightly colored, 
inflorescences larger, with white, open flowers. 

Note. C.multiflora forms hybrids with a number of other species 
of this genus: C.multiflora Bge. XC.insignis A. Pojark. (p. 255); 
C.multiflora Bge.X C. m e 1 a no c ar p a Lodd. (p. 247); C.multiflora 
Bge. X C. r ac em if lor a C.Koch. The latter have been found in the 
Caucasus and occur quite frequently in Central Asia (Dzungarian Ala-Tau. 
Centr. and E. Tien Shan, and Kopet Dagh). They show very varied 
combinations of the parental species characters, being either intermediate 
or else closer to one or the other. In hybrids closer to C.multiflora 
the petioles are rather long; leaves rather sparsely appressed -pubescent 
below, mostly obtuse, obovate; inflorescence loose, profusely branching, 
many -flowered, the pedicels more or less slender and long, usually 
pubescent, sometimes tomentose; fruits mostly glabrous even when young, 
often oblong. In hybrids closer to C.racemi flora the leaves are mostly 
ovate or elliptic, larger than those of C.racemiflo ra (3—3.5 cm long, 
1.5 — 1.8—2 cm broad), with either short or longer petioles, the blade 
rather sparsely pubescent to tomentose below; inflorescences more 
compact, with short thick axes and pedicels, more impoverished, 
corymbiform, usually tomentose-pubescent; fruits hairy when young. 
These hybrids have been described as C.nummularia /3 soongorica 
Rgl. et Herd., Enum. pi. Semen, in Bull. Soc. Nat. Mosc. XXXDC, 2 (1866) 58 
(C.fontanesii7 soongorica Rgl. in A. H. P. II (1873) 313; C. soon - 
gorica M. Pop. in Bull. Soc. Nat. Mosc, nouv. ser.,XLIV (1935) 128). 



Series 2. Insignes A. Pojark.— Fruits black. Inflorescences profusely 
branching, large. Young leaves and inflorescences tomentose-pubescent. 



254 



7. C. insignis A. Pojark. (nom. nov.). — C. num mu lar i a C.Koch, 
Dendrol. (1869) 171 (non Fisch. et Mey., non Lindl.). — C.nummularia 
var. lind 1 ey i Wenz. in Linnaea XXXVIII (1884) 189. — C.arborescens 
Zabel. in Mitteil. d. deutsch. dendr. Gesellsch. (1897) 25 (non Wenz.). - 
C.lindleyi C. K. Schneid., Handb. Laubh. I (1906) 75 7 (non Steud.); 
Fedtsch., Consp. fl. t'urk. Ill (1909) 43; M. Popov in Tr. Prikl. Bot., Gen. i 
SeL.XXII (1929) 447 (ex parte).- Ic. : C.K. Schn., I.e., f.425a-c. 

Tall shrub or small tree; shoots white -tomentose when young, later 
glabrous, red -brown or dark brown; leaves bright green and glabrous 
or scattered -hairy only along the midrib above, initially yellowish- 
tomentose below, later with fine, rather sparse pubescence, broadly obovate 
to orbicular or broadly elliptic, mostly obtuse, often emarginate, less 
often short -pointed and cuneate at base, 2—5 cm long, 1.5—4.8 cm broad, 
on sterile shoots to 6 cm long, 5 cm broad; petioles 4—7 cm long; flowers 
in compound 10— 20 -flowered corymbs, more or less equal to leaves, to 
2—3.5 mm in diameter; inflorescence axes, pedicels, hypanthium, and 
sepals yellowish-tomentose; sepals broadly triangular, erect in fruit, not 
becoming fleshy; petals white; stamens 20; styles 2; fruits globose, 
7— 9 mm in diameter, black with glaucous bloom, open at the apex; nutlets 
1 or 2, large, 5—6 (7) mm long, 4—5 mm broad; style arising from apex of 
nutlet. Fl. June, fr. July. (Plate XIX, Figure 4). 

Among shrubs on stony slopes. — Centr. Asia: T. Sh. (Fergana Range), 
Pam. -Al. (Zeravshan, Gissar, and Darvaz ranges, N. slope of Alai Range). 
Gen. distr. : Iran. — Afghanistan, Ind.-Him. — NW Himalayas. Described 
after a cultivated specimen (from Kashmir ?). Type in Leningrad. 

Economic importance. Deserves attention as an ornamental plant of 
southern districts of the USSR. 

Hybrids of C. in s igni s A. Pojark. X C. mu It if 1 o r a Bge. are 
apparently rather common; leaves are closer toC.multiflora in 
size and shape, but have below the same pubescence as inC.arborescens: 
pubescence of inflorescence varies from slight to dense, persisting until 
fruiting; fruits red, darker than in C. mult if lor a, their apex not entirely 
covered by sepals converging toward the center. Zeravshan and Gissar 
regions. 



Series 3. Racemiflorae A. Pojark. — Fruits red; inflorescences rather 
small, composed of 1—3 (4)corymbs. Inflorescence axes and lower surface 
of leaves white -tomentose. 

8. C. racemiflora (Desf.) C. Koch, Dendrol. I (1869) 170; Fedtsch., Consp. 
Fl. turkest. Ill (1909) 43.- M e s p i 1 u s racemiflora Desf., PL hort. 
Paris, ed. 3 (1829) 409.- C.f ontanesii Spach, Hist. veg. phaner. II (1834) 
77; Tsinzerl. in Izv. Bot. Sada XXIII (1924) 15.- C. torn entosa (non 
Lindl.) Hohen., Enum. pi. Talysch. (1836). — C.nummularia Fisch. et 
Mey., Ind.Sem. Hort.Petr. II (1835) 31; Ldb., Fl. Ross. II, 1, 93; Shmal'g., 
F1.349; M. Popov in Trud. Prikl. Bot., Gen. i Sel. XXII (1929) 446.- 
C.fontanesii adesfontaini et /3 nummularia Rgl.inA.H. P. II 
(1873)312,313; Grossg., Fl. Kavk. IV (1834) 284. - C. r a c em if lo r a a. 



255 



typica (ex parte) et b. num m u 1 a r i a C. K. Schneid., Illustr. Handb. d. 
Laubh. I (1906) 754.- Exs. : Sintenis, Iter transc. -pers. ann. 1900-1901, 
No. 657 (sub C. num m ular ia Fisch. et Mey.). 

Shrub rarely more than 1 m high, erect or often crooked, with curved 
branches, sometimes only 30 cm high, the young shoots densely tomentose, 
one or two-year-old shoots glabrous, dark brown or cherry-red; leaves 
bright green or glaucescent -green, subglabrous to more or less densely 
appressed -hairy above, densely white or yellowish tomentose below, broad 
(width-to-length ratio mostly 1:1, rarely 1 : 2 / 3 ), orbicular, ovate, obovate, 
„ „ or broadly elliptic, obtuse, often emarginate or short -acuminate, often 

mucronulate, 0.5—4 cm long, 0.4—3 cm broad; petioles short, 2—5 mm long, 
tomentose; flowers in erect, 5—9 (12) -flowered, dense compound corymbs 
(half as long as leaves) composed of 2 - or 3 (4)-flowered corymbs or 
umbels, issuing from short (ca. 3 mm long), common inflorescence axis, 
corymbs rarely simple, 3— 4 -flowered; sepals broadly triangular, acute, 
all parts of inflorescence (axes, pedicels, hypanthium, ovary, sepals) 
densely white -tomentose; corolla gaping, ca. 1 cm in diameter, the petals 
spreading, orbicular or obovate; stamens 20; styles 2; fruits 8— 10 mm 
in diameter, broadly ellipsoid to subglobose, glabrous (hairy when young), 
bright red, with usually 2 (rarely 1 ) nutlets; nutlets 5 mm long, 4.5 mm 
broad, flat ventrally and convex dorsally, with short hypostyle and with 
style arising near the apex. Fl. May — June, fr. July — September. 

Shrub thickets on stony slopes. — Caucasus: Dag., W., E . and S. Transc, 
Tal.; Centr. Asia: Dzu.-Tarb., Mtn Turkm. (Kopet Dagh, Balakhan Mountains 
Pam. -Al., T. Sh. Gen. distr. : As. Min., Syria, N. Iran, Him., N. Afr. 
Described after a cultivated specimen of unknown origin from the Paris 
Botanical Garden. Type in Paris; cotype in Leningrad. 

Note. Hybrids of this species are described above: C. r a c e m if 1 o r a 
C.KochX integerrima Med. (p. 248); C.racemiflora C.KochX 
melanocarpa Lodd. (p. 322); C.racemiflora C. Koch. X mult if lo r a 
Bge. (p. 330). 

9. C.taurica A. Pojark. in Notul. syst. VIII, 7-9 (1938).- C. nummularis 
auct. taur. (non Fisch. et Mey.). — C. n um m ul a r ia var. o v a 1 i f o 1 i a (non 
Boiss. ?) Vasil'ev in Tr. Prikl. Bot., Gen. i Sel., ser. VIII, 1 (1932) 400. 

Shrub; young shoots whitish -tomentose, annotinous shoots glabrous, 
dark cherry -red, lustrous; leaves firm but rather thin, glabrous or with 
isolated hairs above, grayish -tomentose below, mostly elliptic, less often 
elongate -ovate (length-to-width ratio 2 :1), obtuse, sometimes slightly 
emarginate or rather short -acuminate, usually with a cartilaginous cusp, 
1—3 cm long, 0.5—1.7 cm broad, rarely to 4.5 cm long and 2.5 cm broad 
on sterile shoots; pedicels tomentose, 3—8 mm long; flowers in erect, 
short (half as long as leaves or less), very compact, usually compound 
corymbs composed of 2 or 3 branchlets bearing (2) 3— 4 -flowered corymbs 
or umbels, less often the flowers in simple corymbs; main inflorescence 
axis 3— 5 mm long, tomentose -pubescent like the secondary axes and the 
very short (l— 2 mm) pedicels; hypanthium, ovary, and subobtuse broadly 
„„„ triangular sepals covered with rather sparse pubescence of appressed 
hairs (not tomentose as in C.racemiflora, and not masking the green 
color); flowers ca. 1 cm in diameter, gaping; petals spreading, orbicular 



256 



or rounded -obovate; stamens 20; styles 2; fruits red, ovoid -globose, 
7—8 mm long, with 2 oblong nutlets 5—6 mm long, 3—3.5 mm broad, flat 
ventrally, convex dorsally, with short hypostyle and with style arising near 
the apex. PL May, fr. July -August. (Plate XIX, Figure 3). 

Shrub thickets on stony mountain slopes.— European part: Crim. 
Endemic. Described from the vicinity of Yalta. Type in Leningrad. 

Note. The Crimean plant is distinguished from C.racemiflora 
C. Koch — widespread in the Caucasus and Central Asia — by its oblong 
leaves and slight pubescence of outer flower parts. 

Economic importance. Ornamental plants, especially in fall during 
fruiting. 



Series 4. Saxatiles A. Pojark. — Fruits black; inflorescence small, 
2—4 -flowered. 

10. C. saxatilis A. Pojark. in Not. syst. Inst. Bot. Ac. Sc. URSS, VIII, 
7-9 (1938). 

Shrub 1.5—1.75 m high; young shoots whitish -tomentose, older shoots 
glabrous, light brown; leaves bright green and glabrous or slightly 
pubescent above, grayish-appressed -tomentose below, elliptic or oval- 
elliptic, less often obovate, mostly rounded -cuneate or broadly cuneate at base, 
obtuse, sometimes emarginate or subacute, mucronate, those on short 
shoots 0.9—2.5 cm long, 0.4—1.8 cm broad, those on long shoots to 4 cm long, 
2.3 cm broad, often ovate, mostly acute; petioles 2.5—4 mm long, tomentose; 
inflorescence shorter than leaves, short -pediceled, in the shape of an 
erect, compound corymb composed of 2— 4 -flowered corymbs; axes and 
pedicels reddish, pubescent or tomentose; flowers unknown; fruits with 
short (1—2 mm) pedicels, obovoid -globose, 6— 8 mm long, 5— 8 mm broad, 
glabrous, dark purple when not fully mature, black with slight glaucous 
bloom when mature; nutlets 2 (l ), obovoid, 4.5—5.5 mm long, 3—4 mm ; 
broad, flat ventrally, convex dorsally, the style arising slightly below the 
apex. Fr. September. 

Among shrubs on stony slopes.— Caucasus: S. and E. Transc. Endemic. 
Described from the vicinity of the village of Chaikend near Kirovabad 
(formerly Gandzha). Type in Leningrad. 

Note. The taxonomic status of this species is not quite clear, since 
its flowers are unknown; according to other characters it is closely 
related to species of the section Chaenopetalum, to which it has been 
provisionally referred. 



334 



Genus 725. CYDONIA * MILL.** 
Mill. Gard. diet. ed. VIII ( 1768) . 

Flowers large, solitary; stamens 20; ovary formed by 5 connate carpels 
(styles 5) wholly sunken in hypanthium, each carpel with numerous, biseriate 
ovules; pome large, without nutlets (endocarp coriaceous), with 15—20 seeds 
in each locule; testa mucilaginous. Leaves simple, entire. One species, 

* This name, used by Hippocrates, may have its origin in the name of the city of Cydon [now Canea] 
on the north coast of Crete or from the semimythical tribe which inhabited that island. 
"" Treatment by A.I. Poyarkova. 



257 



335 



1 . C. oblonga Mill., Gard. diet. ed. VIII (l 768); Popov in Tr. Prikl. Bot., 
Gen.i Sel.XXII, 3 (1929) 434; Grossg., Fl. Kavk. IV (1934) 285.- Pyrus 
cydonia L., Sp. pi. (l 753) 480; M. B., Fl. taur. -cauc. I, 391 . - C y d o n i a 
cydonia Pers., Synops. pi. Ill (1807) 40. — C. v ulg a r i s Pers., 1. c, in 
corrig.; Ldb., Fl. Ross. II, 1 , 101 ; Shmal'g., Fl. I, 351 ; Medv., Der. i kust. 
Kavk. (1919) 132.- Ic: Vol'f and Palib., Opred.der.i kust . (1904), figures 
on pp.469, 470, and 471; Hegi, 111. Fl. v. Mitteleur IV, 2 (1925) f. 1024-1026, 
tab. 145, f.4. 

Small tree or shrub 1.5—5 m high, with thin, scaling bark; shoots 
lanate-tomentose when young, later glabrous; leaves ovate or oval, 
less often orbicular, entire, acute or obtuse, mucronulate, rounded 
at base or slightly cordate, the young leaves arachnoid -tomentose, glabrous 
when fully developed, and dark green above, grayish _ tomentose below, 
to 10 (12) cm long, 7.5 cm broad; stipules glandular-dentate; flowers pale 
pink, to 5 mm in diameter, short -pediceled; pedicels, receptacle, and 
outside of sepals tomentose -pubescent; receptacle ovate; sepals oval, 
glandular -serrate; stamens densely lanate at base and tightly constricted 
by projections of perigynous disk; fruits initially tomentose, glabrous 
at maturity, lemon -colored or dark yellow, sometimes reddening laterally, 
mostly somewhat ribbed, globose (f. maliformis Kirchn., C. ma 1 i f o r m i s 
Mill.) or pyriform (f. pyriformis Kirchn.), in wild quince 2.5—3.5 cm 
long and weighing up to 60—100 g; flesh with numerous grit cells, not very 
succulent, astringent, but very aromatic. Fl. May, fr. September. 
(Plate XXVII, Figure 1 ). 

Forest edges, forest and slopes in the lower mountain belt, not higher 
than 1,400 m; on rather deep, mainly limestone soils.— Caucasus: W. and 
E.Cisc, Dag., E. and S. Transc, Tal. Centr.Asia: Mtn. Turkm. (Kopet Dagh, 
Aidere Gorge). The presence of wild quince in Pam.-Al. has not been 
ascertained; it has been reported for the Bald'zhuan district, at the 
Pulisang Bridge on the Vakhsh River, and for the Zigdi Pass; possibly 
growing wild in W. Tien Shan, near Kara-Ungur (Kattar-Yangak natural 
boundary area). Escaped in Crim. Gen. distr. : Arm. -Kurd. (As. Min.) 
and N. Iran. Widespread in cultivation and naturalized almost throughout 
the Mediterranean area, escaped in many areas of Central Europe. 
Cultivated as far north as Scotland and Norway (63° 51' N.). Described 
from the Danube. Type in London. 

Economic importance. Cultivated for its fruits, which sometimes 
weigh up to 2 kg. Wild quinces bear few fruits (2—10 each) and are not 
an important source of raw material in the national economy. The fruits 
are hardly ever eaten raw but are cooked or baked, used as condiment and 
for the making of jam, jelly, marmalade, syrup and drinks. In medicine 
quince seeds serve as purgatives and laxatives; their mucilage is mainly 
used externally to cover and refresh. The extract prepared from the 
mucilage and iron and an infusion of the latter are used instead of ferric 
malate infusion and extract. The mucilage is used in the textile industry 
for giving a gloss to materials; its aqueous decoction can replace gum 
arabic. Seeds contain 20—22% mucilage and 15% of fatty oils, amygdalin, 
the enzyme emulsin, tannins (in seed coat), proteins, dyes, and 13% ash. 
The colloidal aqueous solution of mucilage (50:1) yields under the action of 
diluted acids 34% of a substance reacting like cellulose. The fat contains 



258 



336 



myrsinic acid. The pulp of the fruit contains 9.6% sugars, mostly levulose, 
malic and tartaric acids, and a small quantity of tannins. Quince is 
also cultivated as an ornamental plant; there is a form with variegated 
leaves (f. marmorata (Dipp.) C. K. Schn.) and a pyramidal form 
(f. pyramidalis (Dipp. ) C . K. Schn. ); tolerant of clipping, used for 
hedgerows; serves as a stock for grafting delicate dwarf varieties 
of pear and apple trees, Eriobotrya japonica and Photinia. 
Requires deep, loose, fertile, moist soil; is cultivated from seeds, coppice 
shoots and root suckers. Provides honey and pollen for bees. The 
whitish wood, regular in structure and medium -hard, is easily cut and 
used for various small articles. Nearly all old trees are affected by 
heartrot caused by the fungus Fomes fulva; another fungus, 
Sclerotinia cydoniae, affects the leaves, decreasing the size of the 
fruits and the general yield. 



Genus 726. PYRUS* L. s. str. ** 

L.Sp.pl.ed.l (1753) 479. 

Trees, less often tall shrubs; leaves with early deciduous stipules, 
alternate, involute in bed, deciduous in winter, entire, less often 
pinnatisect. Flowers in corymbs; calyx deciduous or persistent in fruit. 
Petals white or pinkish, long-clawed. Stamens 20—50. Styles 5, parted 
from the base, glabrous or pubescent. Fruits succulent, pyriform, sometimes 
subglobose, the flesh with more or less abundant grit cells. 

1. All or at least some leaves pinnatisect into narrow lobes 

18. P. regelii Rehd. 

+ All leaves entire 2. 

2. Leaves oval, suborbicular, or broadly elliptic, not more than 2—2.5 
times as long as broad, when fully developed quite glabrous or 
subglabrous 3. 

+ All or most leaves narrowly elliptic or lanceolate, 3 or more times 
as long as broad; if leaves broader, then pubescence persisting 
even on fully developed leaves, or else leaves large, to 8—10 cm long, 
with teeth terminating in a persistent or caducous callous point ... 9. 

3. Leaves acutely dentate, the teeth terminating in a more or less long 
sharp point 4. 

+ Leaves obtusely, less often acutely dentate, but teeth without awnlike 

point 6. 

4. Fruiting pedicels 1.5—2 cm long; leaves 4— 8 cm long 

5. P. ussuriensis Maxim. 

+ Fruiting pedicels 7.5— 8 cm long; leaves to 10—12 cm long 5. 

5. Leaves broadly ovate or suborbicular, with large teeth terminating in a 

a long sharp point. Fruits large, 5—6 cm long 

6. P. asiae -mediae M. Popov. 

+ Leaves ovate or elliptic, with very small, inconspicuously apiculate 

teeth. Fruits smaller, 2— 2.5 cm long 7. P. grossheimii A. Fed. 

6. Fruits small, ca. 1 cm in diameter, with early deciduous scales; 
leaves quite glabrous, coarsely and unequally serrate throughout 
margin 4. P. boissieriana Buhse. 

* From the Roman name of this plant. 
** Treatment by V. P. Maleev. 

259 



337 



338 



Fruits usually larger, with scales persistent or else deciduous when 

fruits mature; leaves entire or serrulate or crenate 7. 

Sepals broadly oval, appressed to fruit; in young plants, all parts 
covered with snow-white tomentose pubescence which later almost 

disappears 3. P. turcomanica Maleev. 

+ Sepals elongate, lanceolate, more or less erect; leaves and other 

parts of plant never with tomentose nor with such a dense pubescence 
8. 

8. Leaves and shoots glabrous; leaves acutely serrate 

2. P. balansae Decaisne. 

+ Young leaves and shoots pubescent; leaves entire, crenate, or serrate, 
but with smaller subobtuse teeth 1 . P. communis L. 

9. Leaves very small, to 3—4 cm long, 1—1.5 cm broad, glabrous, slightly 
pubescent only on the margin 13. P. sosnovskii A. Fed. 

+ Leaves larger 10. 

10. Leaves densely villous or sericeous -pubescent on both sides or at 
least below, the pubescence of long implexed hairs, less often 
pubescence more or less disappearing, but then leaves entire or 
obscurely denticulate only at the apex 11. 

+ Leaves glabrous or subglabrous, sometimes pubescent but the 

pubescence less dense and not composed of implexed hairs; leaves 
conspicuously serrate or crenate, the teeth terminating in callous, 
sometimes early deciduous point 14. 

11. Lateral short shoots thick, coarsely annularly scarred; pubescence 
villous 12. 

+ Lateral short shoots slender; pubescence sericeous 13. 

12. Leaves 3.5—8 cm long, 2—4 cm broad. Tree with more or less 
abundant spines, sometimes with none 8. P. elaeagrifolia Pall. 

+ Leaves longer and narrower, 6—9 cm long, 2—3 cm broad; spines 

absent 9. P. taochia Woron. 

13. Leaves short -petioled or subsessile, longer and narrower, 3—4 times 

as long as broad, entire or obscurely dentate only at the apex 

10. P. salicifolia Pall. 

+ Leaves with petioles to 4 cm long, the blade broader, 1.5—2 times as 

long as broad, conspicuously crenate or serrate t 

11. P. takhtadzhiani A. Fed. 

14. Leaves broadest around the middle or higher 15. 

+ Leaves broadest in lower part 16. 

15. Leaves broadest in the middle, serrulate or obscurely crenate 

12. P. syriaca Boiss. 

+ Leaves broadest above the middle, coarsely and acutely serrate 

14. P. oxyprion Woron. 

16. Leaves densely pubescent below, acutely serrate 

15. P. raddeana Woron. 

+ Leaves less densely pubescent on both sides, crenate -serrate .... 17. 

17. Leaves glabrous, elliptic or broadly elliptic 

16. P. zangezura Maleev. 

+ Leaves usually pubescent, less often glabrous, elongate -lanceolate, 

ligulate 17. P.korshinskyi Litw. 



260 



Series 1. Communes.- Leaves broad, oval or broadly elliptic to suborbic- 
ular, initially more or less pubescent, later glabrous or glabrescent, entire or 
denticulate, the teeth not awned. Calyx persistent in fruit. 

1. P. communis L., Sp. pi. (1753)479; Ldb., Fl. Ross. II, 94; 
Decaisne, I.e., 340; Boiss., Fl. Or. II, 653; Shmal'g., Fl. I, 351 ; C.K.Schn., 
Laubholzk. I (1906) 661; Medvedev, Der. i kust. Kavkaza (1919) 126; 
Voronov in Bullet, of Appl. Bot. a. Plant-Breed. XIV, 3 (1924—1925) 58, 76; 
Popov, I.e., XXII, 3 (1929) 404; Vasil'ev, 1. c, ser. VII, VI (1932) 388.- 
P.achras Gaertn., De fruct. II (1791 ) 44. — P. py r a s t e r Borkh., Handb. 
Forstbot. II (1803) 1287.- Ic: Decaisne, I.e., tab. I; C. K. Schn., 1. c, 
f. 362 f-i, 363 a-e; Vasil'ev, I.e., f. 8; Popov, 1. c., f. 83. 

Tree to 20—30 m high, sometimes a shrub; branches with or without 
spines; buds and shoots glabrous, less often pubescent; petioles about 
as long as the leaf blade, 2—5—7 cm long, initially moJ*e or less pubescent, 
later glabrous; leaves 2—5—7 cm long, 1.5—2.5 cm broad, suborbicular or 
oval, rounded or obscurely cuneate at base, short -tapering, acuminate 
apex, entire or serrulate or crenate on whole or part of margin, initially 
white-arachnoid-pubescent, especially below, later quite glabrous or 
subglabrous, with a stronger pubescence — masking the teeth — only along 
the veins and the leaf margin, lustrous green, lighter below, drying black; 
pedicels 3.5 cm long, pubescent or glabrous; flowers 2.5—3 cm in diameter; 
sepals triangular -lanceolate, densely pubescent like the ovary, erect; 
petals short -clawed, ca. 1.5 cm long, 1 cm broad; fruits pyriform or 
subglobose, very variable in size and shape, to 3—4 cm long, 1.5—2 cm 
broad, green, sometimes reddening, less often yellow. April, May. 

Deciduous, sometimes coniferous forests and shrub thickets, in deep 
soils; sometimes forming pure stands; in the Crimea to 1,100 m, in the 
Caucasus to 2,000 m. — European part : U. Dnp., V.-Don, M. Dnp., L. Don, 
BL, L. V., Crim.; Caucasus: Cisc, Dag., W., E., and S. Transc, Tal.; 
Centr. Asia: apparently only as an escape in Mtn. Turkm., Amu D., 
Fam.-AL, Syr D., T. Sh. Gen. distr.: all of Europe (except Great Britain 
and Denmark and the Iberian Peninsula where it is found only as an 
escape), N.Anatolia, N. Iran. Described from Europe. Type in London. 

Note. Very variable in all its characters — size and shape of 
crown, presence of spines, shape of leaves, degree of pubescence, etc. , 
especially size, shape, and taste of fruit. There are two basic types, differing 
mainly in shape of fruit, with numerous varieties and forms: Achras 
with pyriform and Py raster with globose fruit; these two basic types are 
sometimes described as separate species, but there is no basis for this 
differentiation. Moreover, the cultivated forms derived from P. communis 
without hybridization are often united as var. s ativa DC, distinguished 
by absence of spines and by larger fruits. 

Economic importance. The common pear is the ancestor of up to 1,500 
cultivated forms. The most ancient of these, which have changed little, 
are derived directly from this species; most varieties, however, are the 
result of hybridization with other species. Pear trees were first cultivated 
in ancient Greece, then penetrated into Italy, and later became widespread 
in Europe. Wild pear fruits are usually tart, becoming sweeter if stored. 
They are used in various ways depending on their gustatory qualities: 
they are eaten raw or dried, stewed, or used for making of drinks (kvass, 
cider) or as fodder for cattle. The utilization of wild pear fruits is of 
great importance in Ciscaucasia, where in the Krasnodar Territory 
alone, according to Trusevich, pear forests with an admixture of apple 

261 



340 



trees occupy 30,347 hectares, and the possible yield (together with the 
apples) is estimated at 139 thousand tons. Wild pear fruits contain 
70—85% water, 6—13% sugar, 0.1— 0.2% acids (mainly citric and malic), 
0.29% ash, tannins, etc. The seeds contain 12—21% fatty oils. The wood 
is heavy, specific gravity 0.72, fine-grained, solid, reddish brown, an 
excellent imitation of ebony when covered with black varnish; used for 
lathework, cabinet -making, and musical instruments. A light brown dye 
is obtained from the bark. 

2. P. balansae Decaisne, Le jardin fruitier de Museum I (1871—1872) 
319; Voronov in Bulletin of Appl. Bot.X, IV, 3 (1924-1925) 58, 78.- Ic: 
Decaisne, 1. c, tab. 6. 

Tree unarmed; young shoots and leaves quite glabrous; petioles 
1.2— 3 cm long; leaves elongate -oval, 4— 4.5 cm long, 2— 3 cm broad, 
with acute, tapering apex, with rather large and very acute teeth on the margin; 
fruits pyriform, ca. 2.5 cm long, 2—2.5 cm broad, with pedicels 2.5— 4 cm long. 

Forests and shrub thickets. — Caucasus: W. Transc. (Adzharia). 
Gen. distr. : Lazistan. Described from near the village of Khabakhor in 
Lazistan. Type in Paris. 

Note. Recorded for Adzharia by Yu. N. Voronov; this indication 
must be verified, however, for there are no herbarium specimens of this 
species from Adzharia. 

3. P. turcomanica Maleev in Acta Inst. Bot. Ac. Sc. USSR, ser. I, 3 
(1936) 196.— P. salvifolia Bogushevsky in Bull, of Appl. Botany etc. 
ser. VIII, 1 (1932) 133, non DC- Ic: Bogushevskii, 1. c, f. 57 and 58 (hab.); 
Maleev, 1. c, f. 1. 

Tree to 10— 15 m high, unarmed, with broad, irregular crown; buds and 
other parts of plant densely and softly white -tomentose, later glabrous; 
shoots initially pubescent, later glabrous, initially lustrous red -brown, later 
smooth and gray with small lenticels; petioles 3.5— 5 cm long; leaves 
initially silvery -white below with a very dense pubescence, later glabrescent, 
pubescent only in angles between veins, lustrous green, entire or slightly 
sinuate and obtusely dentate, mainly near the apex, suborbicular or broadly 
oval, 4 — 7 cm long, 3—5 cm broad, rounded or even slightly emarginate at the 
apex, sometimes acutely short -tapering, more or less cuneate at base; 
flowers unknown; pedicels 2—4 cm long, thickening upward; fruits broadly 
pyriform, ca. 2.5 cm broad, 2 cm long, gradually passing into pedicel below, 
flat at the apex, with short, broad, white -pubescent scale lobes spreading 
and appressed to fruit, thick-skinned, with numerous grit cells. April. 

Dry stony slopes, rarely valleys, on deep alluvial soils. — Centr. Asia: 
Mtn. Turkm. (W. Kopet Dagh). Gen. distr. : Iran. (?). Described from 
W. Kopet Dagh. Type in Leningrad. 

Note. Little varying species, clearly differentiated by its snow-white 
tomentose pubescence and very characteristic scale lobes appressed to 
fruit. The aboriginality of this species in W. Kopet Dagh — where it possibly 
represents the remains of an ancient culture — and its distribution outside 
the USSR have still to be ascertained. 

Economic importance. Fruits sweet, with resinous taste, used like those 
of the common wild pear. 



262 



Series 2. Cordatae Maleev. — Proles A r mo r ic an a Decaisne, 1. c, 
122, p.p.— Leaves as in the preceding but quite glabrous and more coarsely 
dentate. Fruits small, with caducous sepals. 

4. P. boissieriana Buhse in Nouv. Me'm. Soc. Nat. Mosc.XII (i860) 87; 
C.K. Schn., Laubholzk. 662; Voronov, 1. c, 58, 78; Popov, 1. c, 405; 
Bogushevskii, 1. c, 128. — P. c o r d at a Decaisne, 1. c, 330 pp.; Boissier, 
1. c, II, 653, p.p.- Ic: C.K. Schn., 1. c, f. 363g, 364 a-b; Bogushevskii, I.e., 
f.461 (hab.). 

Small tree or shrub with irregular crown, unarmed; all parts of plant 
quite glabrous; petioles 2.5—4 cm long, usually longer than, less often 
as long as the blade; leaves lustrous green, drying blackish, suborbicular, 
2—4 cm long and broad, rounded at base, nearly rectilinear or slightly 
emarginate, slightly tapering at the apex, short -acuminate, unequally serrate 
throughout the margin, with subobtuse teeth; pedicels slender, to 4 cm 
long, much longer than fruit; fruits small, ca. 1 cm in diameter, subglobose, 
lustrous, reddish; sepals early caducous. April. (Plate XX, Figure 1 ). 

Dry slopes at 600-1,200 m. - Caucasus: Tal. (?); Centr.Asia: Mtn. 
Turkm. (W. Kopet Dagh, Sumbar and Chandyr river basins). Gen. distr.: 
Iran. (N.). Described from the vicinity of Radkan in the Elburz Range in 
N. Iran. Type in Geneva. 

Economic importance. A species more resistant to drought, valuable 
for hybridization with common pear and as stock for grafting for more 
arid regions; gives rise to abundant root suckers. 



Series 3. Sinenses Maleev.— Proles Mongolica Decaisne, 1. c, 124, 
p. p. — Leaves as in preceding series but acutely dentate, with aristate 
teeth. Scales persistent in fruit. 

5. P. ussuriensis Maxim, in Bull. Acad. Sc. St. Pe'tersb.XV (1857) 135; 
Prim. Florae Amur. (1859) 102; Regel in Gartenfl. X (1864) 374; Rehder 
in Proceed, of the Amer. Acad. 50 (1915) 227; Kom. in A. H. P. XXXIX, 1 
(1923) 75; Skvortsov in Bullet. Jard. Bot. Princ. XXIV (1925) 146; in The 
China Journ.XIV, 6 (1931) 329; Kom. and Alis., Opreditel' II (1932) 637.- 
P. sinensis Lindl. apud Maxim, in Mel. Biolog. (1873) 168 et in Bull, 
de l'Acad. Sc. St. Petersb.XIX (l 874) 172; Korzhinskii in A. H. P. XII 
(1892) 335; Palibin, ibidem XVII, 1 (1899) 75; Kom., ibidem XXII (1904) 476; 
C. K. Schn., Laubholzk. I, 663 p. p. — P. c ommuni s Bunge in Mem. Ac. Sc. 
St. Petersb. II (1833) 10, non L. — P. s inen s is a ussuriensis Makino 
in Tokyo Bot. Mag. XXII (1908) 69.- Ic: Reg., 1. c, tab. 345; Skwortzow 
in The China Journ.XIV, 6 (1931) 330, f. 1-16, tab. 1-3. Vernacular name: 
khingalikhta (Goldi). 

Tree to 10—15 m high, spiny; crown broad, dense; buds and young shoots 
with fugacious lanate pubescence; branchlets gray to brown or nearly black, 
lustrous; petioles 2—6 cm long, initially pubescent, later glabrous; leaves 
suborbicular, sometimes slightly cordate, with long -tapering acute apex, 
densely and acutely serrate with long-apiculate teeth, with pubescence 
fugacious below and persisting longer only on the margin, lustrous green 
above; inflorescence many -flowered; flowers 3— 4 cm in diameter; sepals 



263 



(343) 




PLATE XX. 1 — Pyrus boissieriana Boiss. et Buhse, branchlet with leaves and fruit; 2 — P. uss uri e nsi s 
Maxim., branchlet with leaves and fruit: a) leaf margin; 3 — P. eleagrifolia Pall., branchlet with 
leaves and fruits, a) leaves; 4— P.taochia Woron., branchlet with leaves and fruit; 5 - P. sali c i foli a 
Pall., branchlet with leaves and fruits: a) leaves. 



264 



broadly triangular, tapering at the apex, rather sparsely pubescent above; 
petals ca 2.5 cm long, 1 .5 cm broad, short-clawed, white; fruits short - 
stalked (1 5-2 cm), either dull dingy green or yellowish, often with reddish 

ts ! .5_ 6 .5 cm long, more or less broadly pyriform or subglobose, the 
skin thick, the flesh with very abundant grit cells. May. (Plate XX, Figure 2). 

Forests and river valleys in the lower mountain zone.- Far East: 
Uss. Gen. distr. : Jap. -Ch. (Manchuria, Korea). Described from the Ussun 
River. Type in Leningrad. 

Note This species varies in size, shape, color, and taste of fruit, on 
the basis of which Skvortzov has described several varieties which may 
be of importance for selection. 

Economic importance. The fruits may be eaten raw or dried; to improve 
their taste they are fermented in heaps and then salted. They are sold in 
Manchurian markets as "li." Being very frost -resistant, this species is 
valuable as stock for grafting for northern horticulture and attention is 
being given to this possible use. The dense, heavy wood has the same 
applications as that ofP.commun is. P. us s u r i e n s i s was introduced 
into cultivation in Europe by the Petersburg Botanical Garden from seeds 
collected in the Far East by Maack. 

6. P. asiae -mediae M. Popov in Bull, of Appl. Bot. XXII (1929) 414, 425 
(pro subsp. P. sinensis Lindl.).- Ic: ibidem f. 87. 

Large tree with spreading crown, unarmed; petioles shorter than leaf 
blade 6-8 cm long; leaves large, to 11-12 cm long, 8-9 cm broad, with broad 
rounded or cuneate base, broadly ovate or orbicular, short -tapering and 
acuminate, coarsely serrate with long-awned teeth, dark green and lustrous 
above paler below; flowers small; pedicels long, to 6-7 cm long, as long 
as fru'it: fruits large, 6-7 cm long, yellowish, short -pyriform, watery -sweet, 
. maturing early. May. 

' Apple -and -walnut forests, in river valleys on deep alluvial soil. - Centr. 
Asia: T. Sh. (Pskem River valley). Endemic. Described from Mullyal. 
Type in Leningrad. ,. 

Note. This species has not been fully studied, nor has its distribution 
area been precisely determined; it is closely related to the Chinese 
P lindleyi Rehder (= P. sinensis Lindl.) and to P. o vo i d e a Rehder. 
The calyx persistent in fruit, clearly distinguishes it from P. serotina 
Rehder widely cultivated in E. Asia.. It has not been determined whether 
the Central Asian P.asiae-mediae is wild -growing or an escape. 
The latter is more probable, especially since the varieties cultivated 
and rather widely distributed in Tadzhikistan and Uzbekistan are derived 
from P. s i n e n s i s Lindl. and from its hybrids with F.communisL. 
These varieties are known in Tadzhikistan as "Noshputy" and "Nok ^ 
(see Viktorovskii in "Trudy Tadzhikistanskoi kompleksnoi ekspeditsn 
[Transactions of the Tadzhikistan Combined Expedition], No. XIII, 1935). 

Economic importance . Like the other forms, P. s i n e n s i s s . 1. is 
valuable for cultivation and selection on account of its very sweet fruits, 
much appreciated by the population of Central Asia. In addition, varieties 
of P. sinensis s. 1. and its hybrids with P. c ommuni s are noted for 
their high yields, are highly resistant to fungal diseases, and suffer little 
from Laspeyrisiapyrivora. For these reasons they are particularly 
valuable for cultivation in the southern USSR. 



265 



7. P.grossheimii A. Fed. in Tr. Arm.fil. Akad.Nauk II (1937) 203. 

Tall or medium- sized tree with conical crown and blackish verrucose 
branches; petioles long (to 10 cm); leaves 8 — 10 cm long, 4 — 5 cm broad, 
ovate- elliptic, broadly rounded at base, tapering- acuminate, slightly 
plicate along the midrib, acutely serrulate, lustrous green; flowers unknown; 
fruits with slender pedicels to 5 cm long, globose-pyriform, 2 — 2.5 cm long 
and as broad; sepals spreading, tomentose-pubescent. 

Hornbeam- oak forests (Quercus castaneifolia) of the middle 
forest zone.— Caucasus; Tal. Endemic. Described from near the village 
of Lerik. Type in Erevan. 

Note. According to A. A. Fedorov, this species is distributed "in the 
virgin forests of Talysh." It is interesting to note that pear varieties of 
the series Sinenses are cultivated in Talysh and in Mazanderan; 
however, it still should be determined whether P.grossheimii in 
Talysh is not in fact an escaped form. 



Series 4. Ponticae Maleev. — Proles Pontica Decaisne, 1. c, 123. — 
Leaves narrowly or broadly lanceolate, less often to narrowly oval or 
„ . „ narrowly elliptic, entire or obscurely denticulate, densely villous or 

sericeous-pubescent like other parts of plant, the pubescence persisting at 
least on underside of leaves, less often the pubescence disappearing toward 
fall. Calyx persistent in fruit. 

8. P. elaeagrifolia Pall, in Nova Acta acad. Petrop. VII (1793) 355; 
M.B.,Fl.Taur.-Cauc.l(l808)389; 111(1819)333; Ldb.,Fl.Ross,II,95; Stev.in 
Bull. So c. Nat. Mo sc. XXIX, 170; Decaisne, I.e. ,316; Boiss., I.e., 654; Shmal'g., 
51. 1, 351; Voron.,1. c, 58, 79; Voinov in Zap. Gos.Nikitsk. Op. Bot. Sada 
X,2(1928)55; Vasil'ev, I.e., 392. - P. n i val i s Pall, ex Georgi, Beschr. . 
d. Russ. Reiches (1802) 1014.— P. nivalis var. e 1 a e ag r i f ol i a C.K. 
Schn., Laubholzk. 1 (1906)659.- Ic: Pall., I.e. tab. 10; Decaisne, 1. c. 
tab. 17; C. K. Schn., 1. c. f. 322c, 361k; Vasil'ev., 1. c, f. 11, 12. 

Tree to 10— 15m high, sometimes a spiny, rarely an unarmed shrub; 
buds and young shoots with a very dense grayish- white villous pubescence 
persisting on annotinous shoots and then gradually disappearing; lateral 
reduced shoots very thick, annularly rugose, dark gray, nearly black; 
petioles 3 — 4 cm long; leaves rather variable in shape and size, broadly 
lanceolate, sometimes oboval or subspatulate, usually broadest in upper 
third; less often, mainly at tips of fruiting shoots, leaves broader and 
shorter, subelliptic, more or less rounded or narrowed at base, slightly 
tapering- acuminate, often with a small, as it were fitted mucro, 3.5— 8cm 
long, 2 — 4 cm broad, entire or obscurely dentate at the apex, densely 
grayish- white villous on both sides, the pubescence partly disappearing 
above in fully developed leaf; corymbs many- flowered; pedicels 1.5 — 2 cm 
long, densely villous; flowers smaller than in P. communis; petals 
white, pink tinged, 1 — 1.2 cm long, ca. 0.8 cm broad, with short, pubescent claw; 
sepals densely pubescent, lanceolate, more or less erect in fruit; fruits 
of varying shape from pyriform to flattened- globose, to 3 cm in diameter, 
initially pubescent, yellow- green, sometimes reddening. April — May. 
(Plate XX, Figure 3). 



266 



Dry stony slopes, shrub thickets, pine forests, forest -steppe, also high 
mountains of the Crimean Yaila to 1,100m.— European part: Crim. 
Gen. distr. : Bal. -As. Min. (Anatolia), Arm. -Kurd. (Turkish Armenia). 
Described from the Crimea. Type in London. 

Note. Varies greatly in leaf size and shape and in degree of leaf 
pubescence; this is observed on different trees as well as on different 
shoots of the same tree; size and shape of fruit also very variable. 

Economic importance. Distinguished by its strong drought resistance; 
grows well on dry and stony soils; also very resistant to frost (down to 
minus 20—30°). These properties make it a most valuable species for 
forestry and horticulture, since it can be cultivated in the steppe zone. 
Its importance for dry district horticulture is mainly as a drought -resistant 
stock suited to poor soils, and as such it is widely cultivated in E. Crimea 
(see Voinov, 1. c). This species should also be used for hybridization. 
In addition, it has ornamental value and may be used to provide greenery 
in the same regions. In the Crimea the fruits of P. elaeagrifolia have 
the same uses as those of the common wild pear, since both have the same 
properties. 

P. elaeagrifolia Pall. X P. communis L. — Distinguished from 
P. elaeagrifolia by less dense, more fugacious pubescence of leaves 
and other parts of plants, by subglabrous or quite glabrous branches, 
and by broader leaves resembling those of P. communis, usually 
conspicuously dentate, especially in upper part of blade.— European part: 
Crim. Often in districts where both species occur. 

*9. P.taochia Woron. in Bull. Jard. Bot. Princip. XXVI (1927) 609; 
Bull, of Appl. Bot. XIV, 3 (1924) 82. -P. elaeagrifolia f. taochia Woron. 
apud Diapulis in Fedde, Repert. spec. nov. XXXIV (1933) 37.— Ic. : Voronov 
(1924-1925), I.e., tab. 1. 

Unarmed tree; buds and shoots with a dense grayish white, somewhat 
yellowish villous pubescence; annotinous branchlets with smooth, olive -gray 
bark; lateral shortened shoots very thick, with very coarse annulate scars; 
petioles densely villous, 1—2 cm long; leaves long, elongate -lanceolate or 
narrowly elliptic, relatively longer and narrower than those of P. elaea- 
grifolia, 6—9 cm long, 2—3 cm broad, gradually tapering at base and 
decurrent along petioles or more or less rounded at base, obtuse, with 
small, as it were, fitted mucro, entire or obscurely dentate at the apex, 
densely grayish-villous, the pubescence later partly disappearing 
above; flowers unknown; fruiting pedicels ca. 2.5 cm long; fruits pyriform, 
ca. 1.5 cm in diameter, initially pubescent; sepals narrowly lanceolate, more 
or less erect, covered with dense yellowish -gray pubescence. (Plate XX, 
Figure 4). 

Dry stony sites, often forests and shrub thickets. May occur in central 
Adzharistan (Adzharis-Tskhali River basin). Gen. distr.: Arm. -Kurd, 
(middle reaches of the Chorokh River). Described from the former Artvin 
District (between the villages of Gurdzhan and Khad). Type in Leningrad. 

Note. Distinguished from the closely related P. k ot s c hy an a (Boiss.) 
Decne., from Asia Minor, by its narrower, longer leaves and smaller, 
pyriform fruits. 



267 



J48 10. P.salicifolia Pall., Reise III (1776) Anh. 734; L. fil. Suppl. (1781) 
255; Pall., Fl. Ross. I (1784) 20; Ldb., Fl. Ross. H, 95; Boiss., 1. c, 655; 
Decaisne, 1. c, 310; Shmal'g., 1. c. 352; C. K. Schn., 1. c, 656; Medvedev, 
1. c, 128, Voronov, I.e. (1924-1925) 83; Vinogradov -Nikitin in Bull, of 
Appl. Bot.XXII (1929) 90.- P. eleagrifolia auct. fl. cauc, non Pall. - 
P. ar gy r ophy 11a Diapulis in Notizbl. d. Bot. Gart.u. Mus. Berlin XI (1933) 
884; Fedde, Repert. spec. nov. XXXIV (1933) 37.- Ic: Pall., I.e. (1776) tab. 
N. f. 3; Ej., Fl. Ross. (1784) tab. 9; C. K. Schn., 1. c., f. 360c, d, f. 361c; 
Diapulis in Notizbl. (1933) f. 14. 

Tree to 8—10 m high, sometimes a shrub, profusely branching, with broad 
crown and abundant spines; buds, shoots, and leaves covered with dense, 
grayish -white arachnoid pubescence consisting of sericeous hairs persisting 
on leaves until fall but sometimes — especially above — nearly disappearing; 
lateral reduced shoots slender; leaves subsessile or with petioles short, 
to 1—2 cm, the blade extremely variable in shape from long and narrowly 
lanceolate, 6—9 cm long, 0.5—1 cm broad, to broadly lanceolate, 3—6 cm long, 
1—2 cm broad, broadest usually in the middle, less often higher than the 
middle, gradually tapering at both ends and gradually passing into petiole, 
subobtuse or acuminate, entire, sometimes — mainly on strong vegetative 
shoots — terminal leaves coarsely dentate; flowers in many-flowered 
corymbs; sepals and ovary densely pubescent; sepals more or less 
narrowly long -lanceolate, acuminate, more or less erect in fruit; petals 
with short pubescent claw, 1— 1.3 cm long, 0.5—0.7 cm broad; fruits globose 
or broadly pyriform, 1.5—2 cm long, 1.2—1.8 cm in diameter, pubescent, 
especially in upper part, the pubescence disappearing at maturity of fruit; 
pedicels short, usually shorter than fruit, thickening upward. April. 
(Plate XX, Figure 5). 

Dry stony sites, forest edges, shrub thickets, in the steppe. — Caucasus: 
Cisc. (E.) Dag., E. and S. Transc. Endemic. Described from Ciscaucasia 
from the region between the Kura and Terek rivers, in the "Dubki" Hills 
near the Chervlenaya railroad station. Type in Leningrad. 

Note. Very variable in shape of leaves and shape of fruits, which are 
either subglobose or ovoid -pyriform. The variability in leaf shape can 
be observed even on different shoots of the same tree, which makes it 
impossible to separate any constant forms, at least without detailed 
observation in situ or without special collections. An extremely narrow- 
leaved form from the vicinity of Borzhomi and Tbilisi has been described 
as P. ar gy r ophy 11a Diapulis. Examination of the considerable material 
from these regions has not enabled identification of this species, for it 
falls entirely within the variation range of P. s al i c i f o 1 i a, the narrow - 
leaved form of these regions being linked by a full range of transitions to 
broader -leaved specimens of P. salicifolia growing in the same regions; 
in addition, the narrow -leaved form also occurs in other parts of the 
P.salicifolia distribution area. Broader -leaved specimens of 
P.salicifolia are often confused with P.elaeag rifolia, from which 
they are clearly distinguished by leaf shape and, in particular, by the 
type of pubescence. 

Economic importance. Cultivated as ornamentals. Drought and frost 
resistant plants, undemanding in soil requirements, of great importance 



349 



5773 



268 



in arid regions for hybridization and selection of new species as well as 
for silviculture and as stock for grafting. Developing abundant root suckers 
and with a wide -spreading root system which makes them very valuable 
for fixation of sands. Fruits sweet.but with numerous grit cells, used like 
those of wild P. communis; after fermentation they are sold in the 
markets of Transcaucasia as "panto." 

P. s alic if olia Pall. X P. c o mmun i s L., Shmal'g., 1. c, 351 . — 
Distinguished by its broader leaves with less abundant, fugacious pubescence, 
usually dentate.— Caucasus: occurs in districts where the two species grow 
simultaneously. 

11. P.takhtadzhiani A. Fed. in Tr. Arm. fil. Akad. Nauk II (1937) 208.- 
Ic. : ibid. 

Medium -sized tree; branches with few spines or unarmed, covered 
with dark ash-gray bark; leaves varying in shape — often even on the 
same branch — obovate, rhomboid, or elliptic, acute or obtuse, mostly 
tapering at both ends and acuminate, irregularly serrate or crenate, 
sometimes gray-arachnoid-pubescent below, especially along midrib, 
3— 8 cm long, 2 — 3 cm broad; petioles 2.5—4 cm long, gray -pubescent; 
flowers unknown; fruits large, to 4 cm long, 2.5—3 cm in diameter, obovoid, 
succulent, glabrous, yellow -brown; fruiting pedicels long, as long as or even 
longer than fruit, less often shorter. 

Open juniper and deciduous forests.— Caucasus: S. Transc. (S. Armenia). 
Endemic. Described from near the villages of Khosrov and Garny in 
Armenia. Type in Erevan. 

Note. Closely related to P. c o mmun i s in leaf shape but to 
P. salicifolia in other characters; it is distinguished from the latter 
by leaf shape, long petioles and pedicels, and larger fruits. The possibility 
of its being a hybrid of P. salicifolia X P. communis is, apparently, 
to be excluded, since P. communis does not grow in the distribution 
area of P.takhtadzhiani. 



Series 5. Syriacae Maleev. — Leaves lanceolate to elliptic, when fully 
developed more or less pubescent or glabrous, dentate, the teeth terminating 
in a callous, more or less conspicuous but sometimes early deciduous 
thickening. Calyx persistent in fruit. 

12. P. syriaca Boiss., Diagn. ser. I, X (1849) 1; Fl. Or. II, 655; Decaisne, 
I.e., 320; C.K. Schn.,1. c, 659; Medvedev, 1. c, 12 9; Voronov in Bull, of 
Appl. Bot.XIV, 3 (1924-1925) 83; Diapulis in Fedde, Repert. XXXIV (1933) 
38.- Ic: Decaisne, 1. c, tab. 9; C. K. Schn. f. 361 o - 362 e-f. Diapulis, 
1. c. (1933) tab. CXLI, f. 9-10. 

Tree to 10 m high, with short, thick spines; buds large, 0.6—0.7 cm long, 
broadly oval, with scales dark brown, remote, broad, acuminate, initially 
ciliate -pubescent, soon glabrous; young shoots and petioles with short 
sericeous pubescence disappearing later; branchlets lustrous reddish- 
brown, second -year branchlets covered by uneven gray bark; petioles 
thickish, 1—2.5—5 cm long; leaves initially arachnoid -pubescent, especially 



269 



351 



below, soon quite glabrous, lustrous green, firm, with a very prominent 
network of veins, broadly lanceolate, 3—9 cm long, 2—3 cm broad, broadest 
at or slightly below the middle, gradually tapering toward both ends, rounded 
or slightly cuneate at base, rounded at the apex or acuminate, regularly 
serrulate or nearly crenate, the teeth terminating in a conspicuous callous 
thickening; flowers in few -flowered corymbs to 3 cm in diameter; petals with 
a hairy claw; fruiting pedicels thick, thickening upward, 2—4 cm long; fruits 
pyriform, less often subglobose, 2—2.5 cm long, 1 .5—2 cm in diameter, with 
elongate -lanceolate sepals April. (Plate XXI, Figure l). 

Dry slopes , open forests, shrub thickets. Caucasus: S.Transc. 
Gen. distr. : E. Med. (Cyprus, Syria, E. Anatolia), Arm. -Kurd. (Kurdistan, 
Turkish Armenia). Described from the Anti-Lebanon (Mt. Kassia). 
Type in Geneva. 

Note. In Transcaucasia P. syriaca occurs only in S. A r m e n i a. 
The Caucasian specimens differ somewhat from the Syrian and from the 
Asia Minor specimens in general; Transcaucasian P. syriaca should 
perhaps be separated as a distinct race; lack of material, however, 
makes this impossible at present. 

Economic importance. Rarely in cultivation. May be of value as 
drought resistant stock. In Syria there are cultivated varieties derived 
from P. s y r i a c a, but their fruit is of poorer quality than that of the 
native varieties of P. c o mmun i s. 

13. P. sosnovskii A. Fed. in Sb. Nauchn.tr. Bot. o-va Arm. SSR i Arm. 
fil. Akad. Nauk. I (1938) 3.- Ic: ibidem. 

Small tree or tall shrub, spiny; branches densely leafy, the bark reddish 
or brownish, on perennial fruitbearing branches ash-gray; leaves small, 
to 3 cm long, 1—1.5 cm broad, elliptic or subrhomboid, regularly tapering 
toward both ends, cuneate at base, acuminate, minutely serrulate, glabrous 
or slightly soft -hairy only on the margin, lustrous bright green; petioles 
slender, as long as or somewhat shorter than the blade. Fl. and fr. unknown. 

Juniper stands on stony slopes. — Caucasus: S. Transc. (Garni River 
gorge near Erevan). Endemic. Described from there. Type in Erevan. 

Note. Very distinctive, distinguished from other species of this 
genus by its small leaves and general appearance. 

*14. P. oxyprion Woron. in Bull. Jard. Bot. Princ. XXVI (1927) 608; 
Bull. of Appl. Bot.XIV, 3 (1924-1925) 86.- P. syriaca var. oxyprion 
Diapulis in Fedde, Repert. sp. nov. XXVI (1933) 38.— Ic. : Voronov, 1. c. 
(1924-1925). Plate II. 

Medium-sized tree with broad crown; branches spiny; shoots with 
disappearing arachnoid pubescence; buds dark brown, large, with oval 
or broadly triangular more or less remote scales, with disappearing 
arachnoid pubescence and with short white hairs persisting on back of 
scales; petioles 0.5— 2 cm long; leaves initially arachnoid -pubescent below, 
later glabrous on both sides, firm, lustrous green, irregularly acutely 
serrate, often somewhat curved, oblanceolate, broadest above the middle, 
5—9 cm long, 1—1.5 cm broad, tapering but obtuse at the apex, somewhat 
decurrent along the petioles; flowers unknown; fruiting pedicels 4— 5 cm 



270 



long; fruits pyriform, ca. 2 cm long, 1.5 cm in diameter, pubescent in upper 
part when immature; sepals lanceolate, acute. (Plate XXI, Figure 2). 

Dry slopes; may be found in S. Transc.on the slopes of the Aras River 
valley in its middle reaches. Gen. distr.: Arm. -Kurd. (Turkish Armenia). 
Described from the Agri-Dag Range. Cotype in Leningrad. 

15. P.raddeana Woron. in Bull. Jard. Bot. Princip. XXVI (1927) 608; 
Bull, of Appl. Bot. XIV, 3 (1924-1925) 83.- P. syriaca var.raddeana 
Diapulis in Fedde, Repert. XXXIV (1933) 38. 

Unarmed tree; shoots more or less pubescent or glabrous; buds with 
scales pubescent on the back and on the margin; petioles pubescent, 
3—4 cm long; leaves densely tomentose -pubescent below, initially pubescent, 
later glabrous above, elongate -elliptic, broadest at the middle, gradually 
tapering toward both ends, acute, regularly and acutely serrate -dentate, 
6—8 cm long, 2—4 cm broad. Fl. and fr. unknown. 

Caucasus: S. Transc. (Armenia, Zangezur). Endemic. Described from 
near the village of Lishka. Type in Leningrad. 

Note. A little studied, somewhat dubious species; only sterile branches 
are known; possibly a hybrid of P. s al ic if ol ia X P. sy r ia c a; its 
distribution is also rather uncertain, since Frieck's specimens from the 
vicinity of Kiravokan, reported by Yu. N. Voronov and similar to the type, 
do not correspond to the diagnosis of that species. 

16. P. zangezura Male ev in Acta Inst. Bot. Acad. Sc. URSS, ser. I, f. 3 (1936) 
195. 

Unarmed tree; buds large, ca. 5 cm long, narrowly conical to oval, with 
appressed, broadly oval, acuminate scales persisting in terminal buds, dark 
brown, intially densely pubescent, especially on back of scales, later 
subglabrous; young shoots drying almost black -brown, with fugacious 
pubescent; branchlets covered by gray bark with small lenticels; 
petioles 2— 5 cm long; leaves elliptic, broadly -lanceolate, or elongate -oval, 
broadest in lower part, very large, 5—9 cm long, 3—5 cm broad, broadly 
rounded at base, obtuse or somewhat acuminate, obtusely dentate, almost 
crenate, the teeth terminating in a very small, early deciduous callous 
point, dull green, much paler below, drying black, glabrous when fully 
developed, with scattered villous pubescence only along the veins below; 
flowers unknown; fruits in groups of 7 or 8 on upward -thickening stalks, 
ca.2.5 cm long, obscurely pyriform or subglobose, very hard and stony, 
glabrous; sepals lanate -pubescent (ca. 1.5 cm long in midsummer). 

Upper forest zone. — Caucasus: S. Transc. (Armenia, former Zangezur 
County, near the village of Mazra, in the Sav-Chai River valley, at 1,850 m). 
Endemic. Described from there. Type in Leningrad. 

17. P. korshinskyi Litw. in Travaux du Musee Bot. Ac. Sc. Petersb. I, 
(1902) 17; C.K. Schn., Laubholzk. I, 657; Fedch., Rast. Turkest. (1915) 488; 
Popov, I.e., 401.- Ic: C.K. Schn., I.e., f. 360 g; Popov, 1. c, f. 78. 
Vernacular name: shaking (Tadzhik). 

Medium -sized, unarmed tree; buds large, oval, obtuse, with densely white - 
pubescent oval, somewhat remote scales; young shoots densely white - 
pubescent, annotinous shoots covered with brownish-gray bark; petioles 



271 



353 




PLATE XXI. 1— Pyrus syriaca Boiss.,branchlet with leaves; 2-P.oxyprion Woron.: branchlet 
with leaves and fruits; 3-P.korshi nsky i Litw.: a) leaf; 4-P.regelii Rehd. a) leaf margin; 
5— P.bucharica Litw., branchlet with leaves. 



272 



densely pubsecent, 1.5-5 cm long; leaves linear or elongate -lanceolate, 
ligulate, 5-10 cm long, 2-4 cm broad, broadest in lower part, broadly rounded 
at base or broadly cuneate, gradually long-tapering and acuminate, crenate- 
serrate, the teeth terminating in a subacute callous, later deciduous point, 
more or less densely whitish -pubescent throughout on both sides, when fully 
developed glabrous or subglabrous, lustrous, pubescent only below along the 
veins; corymbs many -flowered; pedicels, ovary, and sepals densely 
pubescent; flowers 2-2.5 cm in diameter; petals elongate -oval, with short, 
glabrous claw; fruiting pedicels 2-2.5 cm long, thickening upward; fruits 
broadly pyriform or subglobose, 3-5 cm long, 3 cm in diameter, the flesh 
with few grit cells; sepals elongate -lanceolate, dentate, erect. May. 
(Plate XXI, Figure 3). 

Dry slopes, shrub thickets, open forests, at 1,200-1,700 m. - Centr. Asia: 
Pam. -AL, T. Sh. (W.). Endemic. Described from the former Andizhan 
County, Fergana Region. Type in Leningrad. 

Note. Varying in shape of leaves and especially in their degree of 
pubescence; fully developed leaves and simultaneously the shoots and 
pedicels either densely pubescent (var.typica M.Popov) or subglabrous 
with traces of pubescence only in upper part of pedicels (var. glabrescens 
M. Popov). 

Economic importance. Most abundant in central and SE Tadzhikistan 
where, according to Linchevskii (Sov'etskaya Botanika [Soviet Botany], No. 1, 
1938) it is widely used by the local population in the planting of nonirrigated 
gardens as drought -resistant stock for cultivated varieties. 

P.korshinskyi Litv. X P. v a v i 1 o v i i M. Popov, 1. c, 403. - Ic. : Popov, 
1. c.,f. 79, 80. 

Leaves oblong -oval or broadly lanceolate, broadly rounded at base, 
gradually tapering at the apex, dentate, glabrous when fully developed; leaf 
shape intermediate between that of the two parent species. Flowers large; 
pedicels lanate. - Centr. Asia: Pam.-Al., T. Sh. (W.). Described from the 
former Osh County. 

Economic importance. The hybrid forms of Central Asia belonging here 
are cultivated and are valuable for selection. 



Series 6. Heterophyllae Maleev.- Leaves quite glabrous like the shoots, 
pinnatisect, sometimes unlobed, dentate, the teeth terminating in a callous 
thickening. Scales persistent in fruit. 

18. P.regelii Rehd. in Journ. Arn. Arb.XX (1939) 97.- P. het e r ophy 11 a 
Rgl. et Schmalh. in A.H. P. V, 2 (1878) 581, non Pott, nee. Steud.; Fedch.,1. c, 
488; Popov, I.e., 399.- Ic: C. K. Schn., 1. c, f. 360 b-g, 361 b; Popov, 1. c, f. 77. 
Vernacular names: Aik-murud (Uzbek), Murudkhyrs (Tadzhik). 

Shrub or small tree to 10 m high, with widely spreading branches and 
abundant long slender spines; buds glabrous, acute, with remote, broadly 
triangular acuminate scales; shoots reddish brown, later gray; petioles 
glabrous, slender, conspicuously thickening at base, long, 2-6 cm; leaves quite 



273 



357 



glabrous, lustrous, light green, thin, to 8—9 cm long, pinnatisect, 
their sessile, more or less broadly lanceolate lobes parted or con- 
fluent, sometimes, in their turn, deeply pinnatisect, acutely serrate, 
acute, 2—5 cm long, 0.3— lcm broad; sometimes some or all leaves 
simple, more or less broadly lanceolate, 5— 7 cm long, 1—2 cm broad; 
pedicels 2— 3 cm long; corymbs many-flowered; flowers 2— 2 .5 cm in 
diameter; petals with a very short claw; calyx and receptacle lanate- 
pubescent, the pubescence more or less disappearing; fruiting sepals 
narrowly long-lanceolate, erect; fruits pyriform, sometimes flattened- 
globose, more or less pubescent below calyx, 2— 3.5cm long and as 
broad. May. (Plate XXI, Figure 4). 

Dry stony slopes, rocks, sometimes in valleys on deeper, moist 
soils; 1, 000-2, 000m. - Centr. Asia: Pam.-AL, T. Sh. (W.). Endemic. 
Described from Kokand and Zeravshan. Type in Leningrad. ningrad. 

Note. Variable in shape of leaves, which sometimes are deeply 
dissected into narrow lobes (most frequently in specimens growing 
on rocks (f. koopmani Spath)); sometimes, in trees growing in 
moister sites and attaining a greater size, most of the leaves are 
simple (f. simplicifolia M.Popov). 

Economic importance. An extremely drought -resistant pear which 
may be widely used for the afforestation of arid regions with poor 
soils. Of interest as drought -resistant stock in grafting. Fruit very 
tart and sticky. According to M.G.Popov, its hybrids were 
important in the selection of cultivated varieties of Central Asian 
pears . 

P . r e gel i Rehd. X P . korshinskyi Litw. — P. bucharica 
Litw., I.e., 18; Fedch., I.e., 488; Popov, I.e., 400. - According to its 
characters, intermediate between the two parent species: leaves 
large, to 13 cm long, 4 cm broad, some simple, others — mostly those 
on vegetative shoots — pinnatisect (var. diversifolia M.Popov), 
less often all leaves simple (var. simplicifolia M.Popov); lobes 
of dissected leaves broader than in P.regelii; simple leaves, 
and lobes of dissected leaves, coarsely and very irregularly duplicato- 
dentate; pubescence more or less strong on buds, shoots, sometimes 
on leaves, which are partly glabrous. (Plate XXI, Figure 5). 

Centr. Asia: Pam.-Al,, T. Sh. (W.). Endemic. Described from 
Darvaz. Type in Leningrad. 

P. regelii Rehd . X P. communis L.— P.vavilovii var . 
glabra M.Popov, I.e., 404.— Little studied form with characters 
intermediate between those of the two parent species. Recorded for 
W. Tien Shan along the Pskem River. 



274 



Genus 72 7. MALUS * MILL.** 

Mill. Gardn. Diet. ed. VIII (1768) 

Medium-sized trees, less often, shrubs; leaves petiolate, with deciduous 
stipules, alternate, revolute in bud, less often plicate, entire or lobed; 
flowers in few-flowered umbellate racemes, usually bisexual; petals with 
a conspicuous claw, white, pink, or red, slightly pubescent; stamens 18 - 50; 
styles more or less connate, glabrous or more or less hairy (usually at 
base); fruit pome, with 2 seeds in each of its 5 locules; testa light brown; 
flesh of fruit in USSR species without grit cells. In the USSR the only 
representatives are of subgenus Eumalus Zabel, characterized by entire 
leaves revolute in bud. 

2 

1 Calyx persistent in fruit 

+ Calyx deciduous in fruit, together with upper part of hypanthium . . 

2 Fruiting sepals free; fruits with a depression at base 3. 

+" Sepals connate at base into tube; fruits with almost rounded base . 

F 8.*M. prunifolia (Willd.) Borkh. 

3. Wild forms (or introduced into cultivation but little changed, not 

hybrids) .' '. \' ' 

+ Cultivated and escaped forms (of hybrid or unknown origin] 

7. *M. domestic a Borkh. 

4. Adult leaves glabrous on both sides '. *. l.M. silvestris Mill. 

+ Adult leaves more or less pubescent below 

5 Shrub form with small flowers (to 3 cm in diameter), native to Mtn. 

Turkm 6. M. turkmenorum Juz. et M. Pop. 

+ Characters not as above " ' 

6. Leaves mostly acutely serrate -dentate (especially in upper part); 

fruits small (European and Caucasian forms) 

+ Leaves mostly crenate- serrate or crenate; fruits often rather large, 
8 to 5 -(7) cm in diameter (Central Asian forms) ... 8. 

7 Leaves mostly with rounded base and conspicuous mucro, usually 
serrulate-dentate throughout the margin, slightly or moderately hairy 
below (South Russian forms) 2 . M. praecox (Pall. ) Borkh. 

+ Leaves usually with cuneate base, muticous or with inconspicuous 

mucro, mostly coarsely serrate- dentate in upper part, mostly with a 
strongly developed tomentose pubescence below (Caucasian forms) 

6 J 3. M. orientalis Uglitz. 

8 Flowers and fruits with or without moderate anthocyan coloring 

4. M. sieversii (Ldb. ) M. Roem. 

+ Flowers and fruits dark" red '.'.'.'.'.'.'.... 5. M. niedzwetzkyana Dieck. 

9. Petioles of adult leaves usually glabrous or subglabrous 10. 

+ Petioles of adult leaves with a more or less dense tomentose 

pubescence ' ' 

10 Leaves finely and acutely serrate-dentate; styles much longer than 

stamens 9. M.baccata (L. ) Borkh. 

Leaves crenate-serrate; styles not longer or very slightly longer 

than stamens 10. M. pallasiana Juz. 



+ 



Roman name for apple tree. 
Treatment by S.V.Yuzepchuk. 



275 



359 



11. Leaves entire at least in lower part, sometimes almost throughout 

their length, obtuse or abruptly mucrolate 

11. M. manshurica (Maxim.) Kom. 

+ Leaves more or less acutely serrate-dentate almost throughout their 

length, with a gradual long sharp point 12. M. sachalinensis Juz. 



Section 1 . PUMILAE Rehder, Man. of cultiv. trees and shrubs N. Amer. 
(1927) 391. 

Series 1. Silvestres Juz.— Fruiting sepals completely free; fruits more 
or less strongly concave at base. 

Note. The taxonomy of representatives of this series has been studied 
very little; there are basic differences of opinion between the various authors 
as to the determination of the number of species and their names. All these 
forms are mostly attributed to one polymorphic species called Pyrus 
malus L. or Malus communis Lam.; at best this is subdivided into 
2 or 3 smaller' species (often the borderlines between them are unclear 
and the names chosen arbitrarily), for example: M. silvestris Mill, and 
M.pumila Mill., or else M. s il ve s t r i s Mill., M. d a sy phy 11 a Borkh., 
and M.pumila Mill, (see especially the last summing-up article by the 
well-known Soviet pomologist V. V. Pashkevich, "Yablonya" [The Apple Tree] 
in the periodical "Priroda" [Nature] 1938, No. 5). The study of this group 
presents tremendous difficulties (at present insuperable), mainly due to: 
(l) the impossibility of differentiating strictly between wild and escaped 
forms and, a fortiori, between their hybrids; (2) the very considerable 
polymorphism of even undoubtedly wild varieties, which makes it difficult to 
apply the geographical-morphological method to this group and to its young, 
still insufficiently stabilized species; (3) the inexactitude and incompleteness 
of herbarium and museum material and of properly performed field 
observations. In accordance with the guidelines adopted in this work, we 
shall still attempt — partly relying on the recently published work of 
G. Koidzumi, "A Synopsis of the genus Malus, " Acta phytot. et geobot. Ill, 
No. 4 (1934) — a slightly different approach to the material at our disposal, 
while specifying that this treatment does not pretend to be definitive, 
attempting only to raise a series of questions for future researchers of this 
group. 

Economic importance. The fruits of the species belonging to this group 
are edible, but those of the wild-growing varieties are seldom eaten raw 
owing to their high acidity; they are usually dried (for stewed fruit, etc.) 
or for the preparation of drinks. Nectariferous. The wood of some forms 
finds applications in carpentry and turnery, but is less valuable than pear- 
wood, because it warps and splits. See also economic importance data cited 
for the individual species. 

1. M. silvestris Mill. Gard., Diet. ed. 8 (1768) No. 1 . - M. ac e r b a 
Merat, Fl. Paris (1812) 187. — M. communis var. sy 1 v e s t r i s Beck, Fl. 
Nied. -Oesterr. (1890) 750. - Py r u s Malus silvestris L., Sp. pi. 
(1753) 479.- P.acerba DC., Prodr. II (l 825) 635.- P. malus var. 
au st era Wallr., Schedae crit. (1822) 215.— P. malus var. glabra 
W. Koch, Syn. Fl. Germ. (1837) 235; Ldb., Fl. Ross. II (1844) 95. - Ic: 



276 



Peterm., Deutschl. Fl. (1849) tab. 26, f. 200; Rchb., Ic, Fl. Germ. XXV (1914) 
111; Hegi, Illustr. Fl. Mittel-Eur. IV, 2, f. 1066, 1067, 1068.- Exs.: 
Woloszczak, Fl. polon. exsicc. No. 947. 

Shrub or tree more than 10 m high with divaricate twigs and dark brown, 
glabrous or initially loosely hairy, more or less spiny branches; entire 
plant subglabrous or slightly hairy (especially on scale margins); leaves 
broadly ovate, broadly elliptic, or suborbicular, mostly rounded or obtuse 
or slightly notched at base, occasionally broadly cuneate, narrowing abruptly 
at the apex to short or shortish, slightly oblique sharp point, serrate - 
dentate or crenate-serrate, often somewhat biserrate-dentate, the teeth 
terminating in a small point (gland), the blade initially covered along the 
veins on both sides with a short crisp, later completely disappearing 
tomentum, conspicuously paler and somewhat lustrous below; petioles 
1 — 3.5 cm long, loosely tomentose, usually glabrous later; flowers in few- 
flowered inflorescences at tips of reduced shoots, to 4 cm in 
diameter; pedicels 1— 2.5 cm long, glabrous or slightly hairy; 
hypanthium glabrous or pubescent at base; sepals 5 — 6 mm long, 
triangular, more or less long -acuminate, glabrous outside, more or 
less tomentose-villous inside; petals 1.3 — 2 cm long, rounded-ovate or 
obovate, abruptly short- clawed, glabrous or slightly pubescent outside, 
white or pink, darker outside; stamens ca. 10 mm long; styles initially 
shorter, later slightly longer than stamens, glabrous, loosely pubescent 
only at base, connate only at the very base; stigma capitate, broader than 
style; fruits 2 — 2.5 cm in diameter, globose or globose- ovoid, yellow- green, 
often slightly reddening on the side facing the sun. May— June. 

Deciduous (especially broadleaf) and mixed (with pine) forests and their 
derivatives. — European part: Lad. -Ilm., U. Dnp., L. Dnp., U. V., V. -Kama; 
reported also for the Caucasus (? ). Gen. distr.: Centr. Eur., Scand., Atl. 
Eur. Described from England. Type unknown. 

Economic importance. Owing to its frost resistance, this species is of 
great importance for the breeding selection of cultivated forms valuable 
for northern regions; for the same reason seedlings of M. silvestris 
are successfully used in these regions as stock for cultivated forms. 

2. M. praecox (Pall. ) Borkh., Handb. Forstb. II (1803) 12 71 saltern 
quoad nomen.- Pyr us praecox Pall., Fl. Ross. I (1784) 22 excl. syn.; 
Ldb., Fl. Ross. II (1844) 97.- P. malus auct. plur. Fl. Ross, saltern p. p. - 
P.malus /3 tomentosa Ldb., I.e., 96 p.p.— Pirus Malus var. 
rossica et var. W o j e j ko w ii Litw. in Maevskii, Fl. Sr. Ross., ed. 5 
(1917) 198.— M. pumila auct. plur. saltern p. p. vix Mill.— M. communis 
Woron. in Acta Hort. B. Ac. Sc. URSS t. XLIII, fasc. 2 (1931) 490. - Ic: 
Woronow, 1. c, f. 419, et 420. 

Shrub or small or medium-sized tree, with rounded crown and slender, 
usually spiny, less often unarmed branches; new annotinous shoots tomentose 
only initially, later glabrous, red-brown; second-year shoots gray-brown, 
with sparse lenticels, very slightly pubescent; leaves 2—10 cm long, 
1 — 4.8 cm broad, broadly ovate to elliptic or obovate, rounded at base, less 
often slightly cuneate or basally notched, obtuse, abruptly tapering to a 
short but conspicuous mucro, mostly acutely serrulate -denticulate or 
biserrulate, less often crenate-serrate, with apiculate teeth, when young 
loosely or sparsely pubescent above, the pubescence later disappearing or 



277 



362 



persisting only along the veins, more densely pubescent below, the finely 
tomentose pubescence persisting, but usually only slightly expressed in 
adult leaves; petioles 0.5— 5 cm long, slender, canaliculate, rather sparsely 
pubescent, often subglabrous on lower side; flowers in few-, usually 3 — 4- 
flowered erect, umbellate inflorescences 4 — 5 cm in diameter, with tomentose- 
villous pedicels 1 — 2 cm long; hypanthium densely white-tomentose; sepals 
triangular-lanceolate, reflexed, acuminate, with glabrous sharp point, 
densely white-tomentose on both sides, less often subglabrous inside; petals 
broadly obovate or ovate, abruptly to short -clawed, white or pink, darker 
outside; styles about as long as stamens or very slightly longer, glabrous or 
tomentose from the base to the middle; stigma capitate, broader than style; 
fruits small, ca. 2 — 2.5 cm in diameter, mostly globose, yellow, with slightly 
tomentose pedicels about as long as the fruit. (Plate XXII, Figure l). 

Broadleaf and mixed forests, forest edges, among shrubs, in S. part along 
river valleys. — European part: U. Dnp., U. V., M. Dnp., V. -Don, V. -Kama, 
Trans v., Bl., L. Don, Crim. ? Described from the banks of the Don and the 
Volga. 

Endemic. Type unkown (possibly in London); cotype (?) in Leningrad. 

Note. The present treatment has given this species a wider scope than 
did Pallas — who considered it primarily as a spiny shrub form from the 
Don and Volga banks — but less wide than that given to it by C. K. Schneider 
(Illustr. Handbuch der Laubholzkunde, Bd. I (1906) 715); the latter author 
included in this species wild Near Asian and Central Asian forms. We have 
followed D. I. Litvinov (in herb.) in referring to M. praecox the tall, wild 
apple trees from the more southerly regions of the European part of the 
USSR, with leaves pubescent below. These forms are sometimes identified 
with M. dasyphylla Borkh., Hand. Forstb. II (1803) 1271, but according to 
C. K.Schneider (l. c, p. 715) the latter is an escaped form of the cultivated 
apple tree, i.e., M. dome s tic a Borkh.; it should be added that all the 
W.European material studied and classified as M. da sy phy 11a Borkh., 
(with one exception) apparently really pertains to the cultivated apple tree. 
The usual identification — originally made by Pallas himself — of 
M. praecox with such cultivated forms as M. pumil a Mill.'s. str. or 
M. frutescens Medik. is apparently without foundation. 

3. M. orientalis Uglitzkich in sched., cfr. Addenda VIII, p. 3 78. - 
M. pumila Grossh., Fl. Kavk. IV (1935) 287 et auct. Fl. cauc.,non Mill. - 
P.malus auct. Fl.cauc— P.malus jbtomentosa Ldb., Fl. Ross. 
II (1844) 96 p.p. 

Medium- sized or more or less tall tree, to 10— 11 m high; branches 
usually unarmed; young shoots dark brown, somewhat tomentose, when 
adult dark gray with sparse lenticels; leaves 3 — 8 cm long, 1.5 — 3.5 cm 
broad, of variable shape, ovate -lanceolate, oblong, short -elliptic, or 
suborbicular, usually cuneately tapering at base, obtuse, less often acuminate 
with inconspicuous or rather short mucro, entire at base, otherwise serrate- 
dentate, less often crenate-serrate, usually with very large acute or subobtuse 
teeth in upper part, thickish; young leaves scattered-hairy above, densely 
white-tomentose below, the adult hairy above only along the veins, otherwise 
quite glabrous, with strongly impressed lateral veins, rather densely, 
sometimes rather sparsely grayish tomentose below, usually very prominently 
veined; petioles 0.5 — 3 cm long, V 4 to % as long as the blade, thickish or 



278 



rather slender, more or less tomentose; flowers 4 — 6 per umbel, ca. 4 cm 
in diameter, with densely tomentose -villous pedicels 8—12 mm long; 
hypanthium obconical, very densely tomentose; sepals rather short, narrowly 
triangular, acute, spreading, densely tomentose outside, subglabrous or 
slightly tomentose inside; petals obovate, narrowing to a conspicuous claw; 
styles about as long as stamens, tomentose at base, otherwise glabrous; 
stigmas clavate, narrow; fruits globose, 2 - 3 cm in diameter, with short, 
densely tomentose pedicels 1-2.5 cm long. April- May. (Plate XXII, 
Figure 2). 

Deciduous (especially broadleaf) mountain forests, forest edges, among 
shrubs, on riverbanks. European part: Crim.? -Caucasus: Cisc, Dag. ?, 
W., E., and S. Transc, Tal. Gen. distr.: As. Min., Iran. Described from the 
Caucasus. Type in Leningrad. 

Note. M. orientalis is described here only provisionally, since it is 
to be described at a later date by Uglitskikh and requires a thorough study 
of the relation of its variation limits. Apparently, it is not the only wild 
Caucasian apple tree. For instance, the interesting Caucasian dwarf apple 
trees originally mainly from Dagestan are not described here due to the 
extreme scarcity of material. 

4. M. sieversii (Ldb.) M.Roera., Syn. Rosifl. (1830) 216.-Pyrus 
sieversii Ldb., Fl. Alt. II (1830) 222; Ldb., Fl. Ross. II (1844) 97. - 
Pyrus n.sp. ? Sievers in Pall. N. nord. Beitr. VII (1796) 292. - P. s ax a - 
til is Schlechtend. in herb. sec. Ldb., Fl. Ross. 1. c, - P. songorica Fisch. 
in herb.- P. malus etMalus pumila auct. fl. As. Med. - Exs.: Kar. et 
Kir. No. 1467. Vernacular name: alma (Kazak). 

Tree 2-10 (14) m high, with gray-brown or dark gray trunk and broad 
crown; branches thick, usually unarmed, less often spiny; (annotinous shoots 
greenish brown, usually more or less anthocy an- colored, more or less 
pubescent; second-year shoots dark gray with remote lenticels; leaves 
large, 6- 11 cm long, 3-5.5 cm broad, short -elliptic to oblong, more or less 
cuneate, less often rounded at base, usually abruptly mucronulate, entire 
at base, otherwise coarsely and shallowly crenate, less often crenate-serrate, 
when adult usually with few hairs only along veins above, with conspicuous 
(prominent in dry leaf) veins, somewhat coriaceous, usually rather strongly, 
sometimes only slightly tomentose throughout; petioles comparatively short 
or rather long, 1.2-3.5 cm, thickish, more or less tomentose; inflorescences 
3 — 5-flowered; flowers large, 3.5— 6 cm in diameter, with long, tomentose - 
villous pedicels; hypanthium densely tomentose -villous; sepals lanceolate, 
finely long-acuminate, reflexed, densely tomentose on both sides; petals 
white-pink; pistils slightly longer than stamens, tomentose to the middle or 
slightly higher; stigma capitate or clavate; fruits rather large (l) 3 — 4 (7) 
cm in diameter, mostly globose or flattened-globose, less often slightly 
elongated, laterally often angular or ribbed, yellow, often partly anthocyan- 
colored; fruiting pedicels mostly long, (l) 2-4 (5) cm, as long as or longer 
than fruit, slightly tomentose-pubescent or subglabrous. April— May. 
(Plate XXII, Figure 3). 

Mountain forests and slopes, mountain stream valleys.— Centr. Asia: 
Dzu.-Tarb., T. Sh., Pam. -Al. Gen. distr.: Dzu. -Kash.? Described from the 
Ul'dzhar River. Type in Leningrad. 



279 



Note. Among the apple trees considered as wild, M. s i e v e r s i i 
undoubtedly is the one with characters closest to those of cultivated forms; 
it is interesting to note that even early authors had doubts concerning its 
"wildness" (Zalesov, for example,judging by the label on one specimen in 
the BIN Herbarium). Contemporary Central Asian botanists and travelers, 
however, insist that M.sieversii is originally from Central Asia. 
3 64 M - G - Popov, however admits the possibility of a widespread hybridization 
between the wild Central Asian apple trees and forms of M. domestica 
Borkh. (oral communication). 

Economic importance. Owing to its comparatively large fruit and 
gustatory qualities, the Central Asian wild apple tree is of particular economic 
interest; in many districts the local population harvests a large yield of 
apples. 

5. M. niedzwetzkyana Dieck, Neuheiten Offerte des Nat. Arb. Zoschen 
(1891) 16; Gard. Chron. 1891, vol. I, 461; Wiener Illustr. Gartenz. (1891) 
164 (ubique sphalm. M e d w i e z t z ky an a); ibidem 1. c, (1892) 18; Koehne 
Deutsch. Dendrol. (1893) 259; Koidz. in Acta Phytotaxon. et Geobat. Ill, No. 4 
(1934) 187.- Pyrus niedzwetzkyana Hemsl. Bot. Magaz. LX (1904) 
tab. 7975.- M.pumila var. n i e d z w e t z ky an a C. K. Schneid., IIL Handb 
Laubh. I (1906) 716 et in Fedde, Repert. Ill (1906) 178; M. Pop. in Bull Appl 
Bot. XXII, No. 3 (1929)432.- Ic: Hemsley, 1. c; Rev. Hort. (1906) 232. 
Vernacular name: kyzyl-alma. 

Small tree; branches unarmed, thickish, dark purple; annotinous shoots 
black; leaves obovate, elliptic, or oblong, tapering toward both ends, often 
cuneate at base, mucronulate, when young loosely tomentose on both sides, 
later only below, rather thick, reddish; petioles long, rather slender or 
thickish, rather densely tomentose like the midrib; flowers intensely purple, 
with slender white -tomentose pedicels; hypanthium and calyx white- 
tomentose on both sides; sepals lanceolate, acuminate; styles slightly 
shorter than stamens, more or less tomentose -lanate at base; pome dark 
violet -red, with pink-purple flesh. Otherwise similar to M. s i e ve r s i i 
(ldb.). M. Roem. 

Mountain forests.- Centr. Asia: T. Sh. Gen. distr.: Dzu. -Kash. (Kuldja, 
Kashgaria). Described after a cultivated specimen grown from seeds of 
a plant found in the Hi District. Type unknown. 

Note. This interesting form should perhaps have been grouped together 
with M.sieversii (Ldb. ) M. Roem., from which it is distinguished only by 
the very strong anthocyan coloration of its parts. In addition, transitional 
forms (or hybrids - M. Popov, 1. c.) may be found between them. 

Economic importance. Cultivated as ornamental and, in its native 
habitat, as a fruit -bearing tree. I. V. Michurin used this form, and especially 
its hybrids, for selection breeding of apple varieties with red-fleshed fruit. 

6. M. turkmenorum Juz. et M. Pop. sp. nova in Addenda VIII, p. 379 
Ic: Bogushevsky in Bull. Appl. Bot. VII ser., No. 1 (1932) 13, f. 10 et 11. 
365 Sh,- ub 2- 3 m high; annotinous shoots anthocyan -colored, strongly 

pubescent; second-year shoots gray, with few lenticels; leaves 6-8 cm 
long, 3.2 - 3.8 cm broad, elliptic, acuminate at both ends, conspicuously veined, 
with broad, acute teeth, strongly pubescent below; petioles 2 -2.5 cm long, 
pubescent; flowers small, ca. 2.5-3 cm in diameter; fruits small, ca. 2 5 cm 



280 



in diameter, globose with broad but shallow depression at base, ribbed, 
yellow; calyx closed in fruit, the sepals long, lanceolate, pubescent; fruiting 
nPdicels drv and long, to 3 cm; seeds flat and long. 

P Mountain slopes and gorges. - Centr. Asia: Mtn. Turkn, (Kopet Dagh). 
Endemic? Described from Mt. Syunt-Dag and the Gyuen Gorge. Type in 

Le Economic importance. The fruits are small and tasteless, hence the tree 
is not valuable for fruit-bearing. However, M. t u r k m e n o r u m is of 
importance as stock for cultivated apple tree varieties, in view of its dwarf 
slz e, drought resistance, and salt tolerance; it has, however, the disadvantage 
of forming coppice shoots. 

7 *M. domestica Borkh., Handb. Forstb. II (1803) 1272 - M. pum ila 
var. domestica C K. Schneid., III. Handb. Laubh. I (l 906 ) 715. - M. d a sy -_ 
phylla var.domestica Koidz. in Acta Phytotax. et Geob. Ill (1 934 1 89. 
Pyrus malus L., Sp. pi. ed. 1 (1753) 479. - P. pu m i 1 a II d o m e s 1 1 c a 
Aschers. et Graebn., Syn. Mitteleur. Fl. VI, 2 (1906) 77. - Ic : Loiseleur- 
Deslongch., Nouv. Duham. VI (1815) 46-55; Schlechtend., Lange u. Schenk, 
Fl. Deutschl. ed. 5, XXV (1886) 2535 (the plate is reproduced \*VarW«h, 
Russk. lekarstv. rast. (1899) Plate 38); Thome, Fl. Deutschl. Ill (1888) 421, 
C. K. Schneid., 1. c, pp. 716, 718. 

Small or often rather large tree with spreading crown, divaricate 
branches, and robust annotinous shoots; leaves mostly large, of variable 
shape, mostly ovate with rounded base, crenate-serrate, with persistent 
pubescence on both sides (much less strong above), short-petioled; flowers 
large, white or pink, usually darker on the outside, with rather short or 
short white-tomentose pedicels; hypanthium and calyx densely tomentose; 
fruits usually large, more than 3 cm in diameter, short-stalked. April 

May. . 

Many varieties cultivated in gardens, often escaping. Gen. distr. : 
now distributed throughout the world. Described after a cultivated specimen. 
Note A large number of cultivated forms derived from various wild 
apple tree species and races have been artificially grouped under the name 
M domestica; it is therefore unavoidable that the morphological 
characteristic of this complex should be rather indistinct. It would have 
366 been more correct to distinguish within the limits of the composite concept 
M domestica a series of quite definite types differing in their origin 
however, a rational classification of cultivated apple trees is entirely a 
thing of the future. Some varieties or groups of varieties have already 
acquired in the literature various binomial designations; such are, for 
instance, M. pumila Mill. Gard., Diet. ed. VIII (1768) s. str., M. f rut esc ens 
Medik., Geschich. d. Bot. (1793) 78, M. a s t r a c a ni c a Dum. -Cours. Bot. 
ed 2 V (1811) 426, M. co st at a Hort., M. c o n o c a r p a Hort, M. striata 
Hortl, M. strepens Hort., M. p r a s o m i 1 a Hort., M. me g am il a Hort., 
and several others. M. dolichomorpha Juz. ined., cultivated in the 
USSR mainly in the Crimea and the Caucasus, should also be mentioned, 
as should the "kandil- sinap" group of varieties characterized, among other 
things, by their pyramidal crown and elongated fruits. 

Economic importance. The fruits are used raw, dried, cooked, canned, 
for the making of jam, marmalade, pastilles, cider, etc. The sap contains 
sugars (glucose, fructose, saccharose), malic acid, tannins and albumins, 



281 



the respective quantities varying from one variety to another. The 
apples also contain a certain amount of vitamins A, B, and C. 



Series 2. Prunifoliae Juz. — Fruiting sepals connate at base into tube, 
acuminate; fruits with almost rounded base. 

8. *M. prunifolia ( Willd.) Borkh., Handb. Forstb. II (1803) 127 8.- 
M. hybrida Lois. -Deslongch. in Nouv. Duham. VI (1815) 140.— Py r u s 
prunifolia Willd., Phytogr. I (1794) 8.— Ic: Lois. -Deslongch., 1. c, 
tab. 42, f. 1; Bot. Mag. tab. 6158; C. K. Schneid., III. Handb. Laubh. I (1906) 
718,720. Vernacular names: yablonya kitaiskaya [Chinesel, kitaika. 

Small tree to 10 m high, the branches pubescent when young; leaves ovate 
or elliptic, abruptly acuminate, regularly and acutely serrulate, initially 
scattered -hairy below, later glabrous or pubescent only along the veins with 
slender petioles V3 as long as or equal to the blade; flowers white; sepals 
lanceolate, divergent, glabrous or tomentose inside, sometimes also pubescent 
outside but less densely so, persistent in fruit, but connate at base into short 
„ Q tube; petals short-clawed; fruits to 2 cm in diameter, globose or ovoid, very 
slightly concave at base, yellow or red. 

Cultivated in gardens. Gen. distr. : known beyond all doubt only in 
cultivation (believed to be native to N. China). Described after a cultivated 
specimen. Type in Berlin. 

Note. A hybrid origin (M. pumil a Mill X M. b a c c a t a (L. ) Borkh.) 
is often attributed to this plant; this opinion has, however, been disputed 
(Rehder, Asami); compare C. G. Dahl., M a 1 u s prunifolia Borkh. etc., 
Svensk. Bot. Tidskr. Bd. 30, H. 3 (1936) 483 — 392. Varies mainly in size and 
color of fruit. — Apart from the true M. prunifolia, the closely related 
M. cerasi folia Spach, Hist. Veg. II (1834) 152, is also sometimes cultivated 
in gardens. It differs from the former mainly in its partly or 
completely caducous fruiting calyx, and is usually taken for a hybrid of 
M. prunifolia X M.baccata. 

Economic importance. Cultivated as ornamental; the fruits are known 
in the Soviet Union as "Chinese apples" or "paradise apple" and are used 
for jam. This species and its hybrids have acquired a considerable 
importance for selection breeding especially that of frost-resistant varieties. 
Especially significant results were achieved by M. I. Michurin, who succeeded 
in creating a number of most valuable varieties suited to the Far North 
by hybridization of different varieties of M.domestica with M. pruni- 
folia. 



Section 2. BACCATAE Rehder, Man. of Cultiv. Trees and Shrubs 
N. Amer. (1927) 391. 

9. *M. baccata (L.) Borkh., Handb. Forst. Bot. II (1803) 1290.- Pyrus 
baccata L., Mant. pi. I (1767) 7 5 excl. syn. Ammani et loc. nat. -P.micro- 
c ar pa Wendl. ex C. Koch, Dendrol. I (1869) 211.— Pyrus vel M. s inens is 
hort. fide C. K. Schneid., III. Handb. Laubh. II (1906) 721.- ? P. baccata 
var. a) genuina, 8) praecox, 7) aurantiaca, 6) latifolia Rgl. in 
Gartenfl. (1862) 202. -Ic: J. Kern., Abbild. Oekon. Pfl. VIII (1796) 756; 
C. K. Schn., III. Handb. I (1906) 718, 720. 



282 



nz t ( 




PLATE XXII. 1-Malus praeeox (Pall.) Borkh., branchlct with flowers: a) leaf, b 1 Style, c) fruit; 
2-M.orinct a 1 i s Uglltz., flower: a) leaf, b) fruit; 3— M.sieversil (Ldb.) M.Roem., branchlct with 
fruits (immature): a) leaf; 4-M.pa 11 asi ana Juz., branchlct with flowers: a) leaf, h) fruits; 
5-M.manshurica (Maxim.) Kom.,branchlet with fruits: a) leaf; G — M.sa eha 1 i n e n si s Juz, 
branchlet with fruits: a) leaf. 



283 



Small tree to 10m high; young branches glabrous; leaves to 6cm long, 
elliptic, ovate, or oblong, rounded or somewhat narrowed at base, acute or 
abruptly acuminate, mucronulate, acutely serrate-dentate, glabrous or 
Initially pubescent along the veins, with Long, glabrous petioles; stipules 
glabrous; flowers few [to 8) in each umbellate inflorescence; pedicels 
[especially Ln fruit) shorter than in the following species, 1 —3 cm long, 

ib Sj sepals twice as I as hypanthium, lanceolate, glabrous outside 

Like the hypanthium, tomentosi inside; petals obovate or oblong, glabrous; 
styles much Longer than stamens, glabrous or lanate at base; fruits globose, 
to i ■ in Long, somewhat depressed, concave at base and apex, yellow, 
redde g in the sun. 

Cultivated Ln gardens. Gen.distr.: Jap. -Ch. ? Described after a 
cultivated specimen. Type in I iondon. 

N o I. e. < '• I • I Ichneider, L. c, 721 , has drawn attention to the differences 
between the typical LVL- bac c ata, known in cultivation, and the Siberian forms 
which even Linnaeus considered as belonging here. 

Economic importance. Ornamental. Like the following species 
Lmportanl for selection breeding (by crossing with cultivated varieties) of 
particular frosl resistant varieties with small fruit called "crab-apples"; 
these a ri ■ n< it very tasty. 

io. M. pallasiana Juz. nom. nov. Pyrus baccata Pall., Fi. Ross, 
t. i, p. I (1784) 23; Ldb., Fl. Ross. II (1844), non L. s. str. — P. b ac cat a 
var. sibirica Maxim, in Mel. biol. IX (1873) 166; Bull. Ac. Petersb. XIX 
(1874) L70. M;il ii.". baccata var.-a. sibirica C. K. Schneid., Ill 
Handb. Laubh. 1 (I 906) 720. IVI. sibirica Kom. in Kom. et Klobuk. -Mis., 
Key for the PI. of the Par Kast. Reg. TI (1932) 638, non Borkh.- Ic: Pall., 
L.c.,tab.X. Exs.: HFRNo.1162. Vernacular name: yablonya sibirskaya 
[Siberian!. 

SmalJ tree with short, mostly floxuous trunk and rounded crown; bark 
rugose, scabrous, ash gray; branch.':, virgate, red-brown, glabrous; leaves 
2.5 8cm Long, L.3 5.5cm broad, ovate or short-elliptic, rounded or 
narrowed at base, abruptly mucronate, obtusely crenate-serrate throughout 
the margin, slightly pubescent when young along the midrib above, later 
usually glabrous on both sides, the petiole rather long, 1.2 —4 cm, somewhat 
pubescent, usually quite glabrous near the tip; stipules narrowly linear, 
glabrous; flowers 2 4 cm in diameter, 4 — 8 in each umbel, with very 
Long, slender, glabrous pedicels 1.5 — 4.5 cm long; hypanthium tubular- 
campanulate, constricted in upper part, glabrous; sepals spreading, linear- 
Lanceolate, acute, more or Less tomentose, usually only inside, sometimes on 
both sales.; petals ol 1 1 < >ng <>v ale, tapering at base to a short claw, white; 
stamens ca. 20, Less than half as long as petals; styles 0.5 — 1 cm long, 
as Long as or verj [lightly longer than stamens, villous at base, occasionally 
glabrous (var. I e i os t y I a Kupr. ); fruits small, to 1 cm long, globose, 
Mow with reddish tinge, Ions, pedieeled. (Plate XXII, Figure 4). 

Forests, forest edges and shrubs, mainly in valleys and along riverbanks — 
E.Siberia: Ang. -Say., Dau.; Far East: Ze. -Bu., Uda, Uss. Gen. distr.: 
Jap. i h., Mono. Described from Transbaikalia. Type in Leningrad. 

N ot e. Following V. L. Komarov, we consider this plant as a separate 
species distinct from the true M. baccata (L.) Borkh. known only in 
cultivation. However, the name M. sibirica (Maxim.) Kom. used by 



284 



371 



372 



V. L. Komarov cannot be retained owing to the existence of an earlier 
homonym IVI. sibir i c a Borkh., attributed by C. K. Schneider (with a 
question mark) to the synonyms of M. as tr ac anic a Dum. Cours. (see 
note to IVI . <1 o m e s t i c a). 

Economic importance. As for the preceding species. 

11. M.manshurica (Maxim.) Kom. in Tr. pochv. -bot. eksp. issled. Az* 
Rossii, Vol. II, 191 3, No. 2, 1917) 93, iiom.j Skvortzow in Bull. Jard. Bot. Rep. 
Russe, t. XXIV (1925) 146.- Pyrus baccata var. manshur ic a Maxim, 
in Mel. biol. IX (1873) 166; liulJ. Ac-.. I. iV-t.-rsi,. i ' I :;v 1 ; ivo. Mains 
baccata var. m ansh ur i ca C. K. Schneid., 111. Handb. Laubh. I (1906) 721 
6t in Fedde, Repert. Ill (1906) 178.- Mains baccata var. manshur ica 
r. Latif oli a Koidz., Consp. Rosac. Jap. (J 91 3) 84. M. m a a sh u rica 
var.genuina et var. Gor d e i e v i Skvortzow, 1. c, Ic: C. K. Schneid., 

in. i ia i kDi. i ,auhh. I (J !)07j 718; Likchonos, Selektsia yabloni [Apple Selection] 
(1936) 134, f. 26. 

Tree to 30 m high with black bark; young branches reddish yellow, 
glabrous; leaves broadly oval, ovate, or obovate, usually rounded at base, 
less often somewhat narrowed, abruptly short-acuminate, usually crenulate 
dentate, less often irregularly denticulate, especially in upper half, subentii ■ 
in lower half, sometimes throughout margin, pubescent when .young, 
especially below, when adult pubescent only along the veins below, Less often 
finely tomentose throughout lower surface; petioles long, 1 -4cm, finely 
tomentose; stipules mostly pubescent; flowers 3 8 in each umbellate 
inflorescence at tips of reduced shoots, 3 — 4 cm in diameter, aromatic; 
pedicels 2.5 -4 cm Jong, pubescent or glabrous; hypanthium tomentose or 
glabrous; sepals lanceolate, acuminate, tomentose on both sides or only 
Lnside; petals 18-22 mm long, oblong, white, often slightly pink outside; 
stamens 18-22; styles 8-8.5 mm long, as long as stamens, villous at base; 
fruits 9 — 15mm long, 9 — 12 mm in diameter, usually oblong, less often 
globose. (Plate XXII, figure 5). 

Riverbanks.- Far East: Uss. Gen. distr. : Jap. -Ch. (N. Manchuria). 
Described from Pos'et Bay and other sites in Manchuria and the Ussuri 
region. Type and paratype in Leningrad. 

12. M. sachalinensis Juz. nov. spec, in Addenda VIII, p. 493. ivi. baccata 
var. s achalinen s i s Kom. in sched. M.manshurica Koidz. in Acta 
Phyt. et Geob. Ill (1.934) 182 p. p. (quoad pi. sach. ). 

Shrub, similar to IVI. m a n s h u r j c a, from which it is distinguished by its 
gradually long-acuminate leaves, subacutely or acutely serrate -dentate 
almost throughout their length, coarsely, acutely, and finely duplicato- 
dentate toward the apex; petioles less density but still conspicuously 
pubescent; pedicels glabrous; hypanthium and calyx mostly glabrous 
outside, less often rather sparsely tomentose; fruits globose or slightly 
oblong. June. (Plate XXII, Figure 6). 

River valleys.— Far East: Sakh. Endemic. Described from Sakhalin. 
Type m Len ingrad. 



2X5 



Genus 728. SORBUS L. 

L. Sp. pi. (1753) 477. 

Flowers in more or less many-flowered, corymbiform inflorescences. 
Hypanthium urn-shaped; sepals 5, triangular; petals orbicular or ovate, 
with or without short claw, white or yellowish white, less often pink. Stamens 
15 — 25. Carpels 2 — 5, adnate by the back to the urn-shaped hypanthium, 
each carpel with 2 ovules, one of which usually does not reach maturity. 
Stigma flat, not broader than style. Fruit globose or ovoid, red, yellow, or 
light brown, the cell walls hard or membranous. Seeds oblong, triangular, 
acute at both ends. Trees or shrubs, mostly with deciduous stipulate leaves, 
widespread in the Temperate Zone of the northern hemisphere. Type species 
of the genus is S. aucuparia or S. domestic a. 

1. Leaves imparipinnate, with serrate leaflets 

Subgenus 1. Eu-Sorbus Kom. 

+ Leave simple, entire or pinnatisect, sometimes pinnate only at 

base Subgenus 2.Hahnia Medik. (see p. 296). 

Subgenus 1. Eu-Sorbus Kom.*— Leaves imparipinnate, with serrate leaflets. 
Carpels mostly adnate to hypanthium only to lz> usually connate only in 
lower part, free in upper part, less often completely connate. Styles 2 — 5 
(mostly 3), usually free. Cell walls membranous, less often rather hard. 

1. Styles 5; fruits yellow, pyriform 1 . S. domestica L. 

+ Styles 2—4; fruits usually red, less often yellow, ovoid or globose . 

2. 

„„„ 2. Shrubs; leaves dark green above, lustrous, as if slightly varnished 

+ Trees sometimes accidentally shrublike (always shrubs in the Far 

North); leaves grayish green and dull above 4. 

3. Leaves 4- or 5-paired; petals glabrous above 

2. S. sambucifolia Trautv. 

+ Leaves 5- or 6 -paired; petals with lanate beards in the middle of . 

upper surface 3. S. schneideriana Koehne. 

4. Leaves glabrous below, with no hairs even along the veins; 
inflorescence branches also glabrous 5. 

+ Leaves more or less hairy below, when old hairy only along the 

midrib 13. 

5. Inflorescence impoverished, less than 5 cm in diameter 6. 

+ Inflorescence developed, considerably more than 5 (mostly more 

than 10) cm in diameter 7. 

6. Sepals with few hairs or cilia 6.polaris Koehne. 

+ Sepals without hairs or cilia 12. S. anadyrensis Kom. 

7. Leaves devoid of pubescence but grayish owing to their peculiar 
epidermal structure 8. 

+ Leaves pale green or light green below 11. 



Treatment by V.L.Komarov. 



286 



8. Sepals glandular-ciliate on the margin, the base plumose, often 

glabrescent later (smooth variety of common Sorbus) 

S. aucuparia var. glabra Gilib. 

+ Sepals without hairs or with simple unicellular hairs 9. 

9. Some stipules not deciduous, broad, with large teeth, leaflets often 
entire from base with teeth in upper part .... 8. S. amurensis Koehne. 

+ Stipules usually all deciduous; leaflets more regularly serrate • ■ • 

10. 

10. Leaflets rather narrow, seldom more than 1.5 cm broad 

7. S. sibirica Hedl. 

+ Leaflets rather broad, more than 2 cm broad and firmer . 

9. S. Boissieri C.K. Schn. 

11. Flowers to 2 cm in diameter; inflorescence branches red; fruits 

to 12 mm in diameter 13. S. tianshanica Rupr. 

+• Flowers ca. 1 cm in diameter; inflorescence branches only 

occasionally reddening; fruits less than 1 cm in diameter 12. 

12. Inflorescence spreading; leaves longer, the leaflets acutely biserrate, 
their teeth longer, aristate 10. S. commixta Hedl. 

+ Inflorescence more condensed; leaves shorter, the leaflets serrate 

with large teeth U.S. kamtschatcensis Kom. 

13. Leaflets thin, usually short-acuminate or even subobtuse, plumose - 
pubescent below; common petiole and inflorescence axes densely 
lanate, the axes spreading; inflorescence ca.lOcm and flowers 
ca.lOmm in diameter; calyx teeth short, deltoid; fruits globose, only 
slightly longer or shorter than their diameter 4. S. aucuparia L. 

+ Leaves firm, often mucronate, with scattered lanate hairs on both 

sides; common petiole and erect inflorescence branches glabrous or 
very slightly pubescent; inflorescence 6 — 8 cm and flower to 12 mm 
in diameter; calyx teeth with convex sides above; fruits usually 
longer than broad 5. S. glabrata Hedl. 

Note. Not included in the hybrid table between S. auc u p ar i a and 
S. f e n n i c a. The hybrid is easily distinguished by its very large terminal leaflet. 
It is indicated for SW Finland, and occurs in the Soviet Union only in gardens. 
It also bears the species name S.meinichii Hedl. 



Section 1. CORMUS Spach, Hist. Ve'g. Phan. II (1834) 96, pro gen. - 
Ovary of 5 connate carpels; styles 5, free or slightly connate, hairy at base; 
petals with beards at base; fruits large, 1.5 — 3 cm long. 

1. S. domestica L., Sp. pi. (1753) 477; C. K. Schn., Ill Laubholzk. I, 683; 
Hayek, Prodr. Fl. Bale I, 747; Shmal'g., Fl. I, 353; K. Zelenetsk., Fl. Kryma 
(1905), 253; Medved., Der. i kust. Kavk. (l 91 9) 120; Grossg., Fl. Kavk. 
IV, 287.- Pirus domestica Sm., Engl. bot. V (1796) 350. — C o rm u s 
domestica Spach, 1. c, 97. — M e s p i lu s domestica AIL, Fl. Pedem. 
II (1785) 141.— Pyrus sorbus Gaertn., De fruct. et semin. II (1788) 
43, tab. 87. — Fench; sorbier, cornier; German: Seierling; Serbian: 
oskwausha. 



287 



375 



Tree 4 — 10 m high; bark scabrous, gray, on young branches olive -gray 
or red-brown: buds glabrous, glutinous, somewhat pubescent at the ends; 
leaves 7 — 10 -paired, to 20 cm long, lanceolate, acute; leaflets broad, 
3 — 50 mm long, 15 — 2 mm broad, or else narrow, 40 — 50 mm long, 15 — 20 mm 
broad, acutely serrate, smooth above, deep green, initially tomentose like 
the petioles, loosing their pubescence at maturity; inflorescence branching, 
densely tomentose, broadly pyramidal, 6 — 10 cm diameter; flowers 1.5 cm 
in diameter; fruits globose (var. pomifera Hayne) or pyriform (var. 
pyrifera Hayne), greenish yellow or red, often brownish red; seeds 
flattened, with subacute margin. Fl. May, fr. September — October. 

Deciduous forests on mountain slopes.— European part: Crim. (southern 
slopes of the Crimean Mountains); Caucasus: Cisc. (until now doubtful; 
most probably in mountains near Novorossiisk). Gen. distr. : mountains of 
Bal. -As. Min., Med., S. part of Centr. Eur. Cultivated in gardens of Central 
and Atlantic Europe and sometimes escaping. 

Note. Wood fine-grained, hard, heavy, reddish white, used for furniture, 
lathework, and small carved articles. The fruit is used for kvass (a fermented 
drink), jam, etc. According toHegi, one tree yields from 6 to 10 centners* of 
fruit. 

Section 2. AUCUPARIA Medik. Phil. Bot. I (l 789) 138.- Ovary of 2-4, 
rarely 5 carpels connate below, free in upper part; styles 2—4, rarely 5, 
free; fruits small, usually less than 1.5 cm in diameter. 



Series 1 . Lucidae Kom. — Shrubs; leaflets lustrous above, dull below, 
smooth; winter buds glutinous, smooth or slightly rufous -pubescent. 

2. S. sambucifolia Roem., Syn. Mon. Ill (1847) 13 9; Hedlund in Sw. Vet. 
Akad. Handl. 35 (1901) No. 1, 37; Kom., Fl. Kamch. II, 237; Hulten, Fl. 
Kamtch. II, 46. — Py r us (s or bu s) sambucifolia Cham, et Schlecht. 
in Linnaea II (1827) 36 et in Linnaea VI (1831) 590; Ldb., Fl. Ross. II, 
99.— Ic: Kom., 1. c, tab. XVIII; C. K. Schn., III. Laubholzk. I, 668. Vernacular 
name: ryabinnik. 

Shrub to 1 — 2 m high; young branches erect, dark brown with glaucous 
bloom, the old branches yellowish gray or gray with very conspicuous 
lenticels; buds acute, more or less glutinous, glabrous, less often slightly 
pubescent, the scales sometimes ciliate-margined; leaves pinnate, with 
7 — 15 oval or oval-lanceolate, acute or acuminate leaflets, dark green and 
lustrous above, pale below, coarsely serrate, often with soft white hairs as 
on the veins; the blade 10 — 18 cm long, 6 — 12 cm broad, the leaflets 3 — 8 cm 
long, 1 —2.5 cm broad; stipules rufous -pubescent, lanceolate, early 
deciduous; common petiole reddish; inflorescence compound corymb 
5 — 10 cm in diameter, its branches and pedicels with sparse rufous hairs; 
flowers ca. 12 mm in diameter, reddish or white; sepals deltoid, straight, 
entire, slightly pubescent only on the margin; petals reddish or white, 
glabrous above; ovary rather sparsely tomentose, gray; bracts early 
deciduous, narrowly linear, ferruginous, lanate; fruits succulent, bright red, 
ellipsoid, globose, or oblong, acid but pleasant to the taste, extremely 



[Metric centner = 100 kg.] 



288 



abundant; in immature fruit the divergent calyx teeth forming a rather 
large crown. Fl. June — July, fr. September — October. 

Growing in large masses in undergrowth ofBetula ermani forests, 
also together with alder and Siberian stone pine stands in subalpine shaul 
thickets near the upper timberline, occasionally forming pure stands; in 
addition, occurs at forest edges and in separate groups; always on dry sandy 
or stony soils.— Arctic: An.; Far East: Kamch., Okh., Sakh. Gen. distr. : 
Kurile Islands, S. Sakh. Described from Petropavlovsk-Kamchatskii. 

Economic importance. Shelter and fodder for cattle. The fruits make 
good jams. 

3. S. schneideriana Koehne in Fedde, Repert. X (1912) 503; Kom. and 
Alis., Opred. II, 637.— Pyrus sambucif olia Maxim. Prim., Fl. Am. (1859) 
(1859) 103 (non Cham, et Schltd.). — Local names: Mis'la, Mirentschura, 
Mirengkola. 

Shrub 1— 2.5 m high; branches 2 mm in diameter; stipules small, 
lanceolate, with rufous hairs, with few long brown-black cilia on the margin; 
leaves 11.5 — 14cm long, 5- or 6-paired; leaflets oblong, short- acuminate, 
entire in lower part, doubly incised-serrate higher, 3.5 — 4.5 cm long; 
inflorescence ca.8cm in diameter, rather loose, smooth, its lower branch 
remote; sepals ciliate; petals barbate above, as long as stamens; carpels 
tomentose at the apex; pistils hairy at base; fruits bright red, ovoid - 
globose. Fl. June, fr. September. (Plate XXIII, Figure 2). 

Woody mountain slopes, mainly along banks of forest brooks, forest 
edges, often together with alders. — Far East: Uss. — in the S., at upper 
timberline below balds. Gen. distr. : Jap. -Ch. (Korea). Described from 
the lower reaches of the Amur. 



Series 2. Aucupariae Kom.— Winter buds with a more or less dense 
white sericeous pubescence or glabrous but not glutinous; petioles of lower 
leaflets 25 — 40mm long; leaves 4 — 8-paired; leaflets 3 times as long as 
broad; stamens as long as petals; carpels quite free in upper part; fruits 
slightly concave above, the fleshy sepals horizontally involute. 

4. S. aucuparia L., Sp. pi. (1753); Hedlund in Sv. Vet. Akad. Handl. 35, I, 
46; Shmal'g., Fl. I, 353; Mela Su omen Kasvio 358; Perfil'ev, FL Sev. Kraya 
II, 173; Rollov, Dikor. r. Kavkaza, 481; Grossgeim, Fl. Kavkaza IV, 287; 
Ldb., Fl. Ross. II, 100. — Py ru s aucuparia Gaertn., de' fruct. et semin. 
II (1788) 45, tab. 87. Finnish name: Kotipihlaj. 

Tree 4 — 15 m, even to 20 m high, with gray, smooth bark and pubescent 
young branches; buds tomentose; leaves 4 — 7 -paired, the leaflets oblong 
or oblong-lanceolate, entire in lower part, serrate higher, mat-green above, 
glaucous or grayish below, more or less hairy (sometimes quite glabrous 
and firmer, var. glabra Gilib., Fl. lithuan. V (l 782) 233, pro sp.), 
inflorescences pubescent, less often inflorescence branches glabrous, 
5 — 10 cm in diameter; flowers 0.8 — 1.5 cm in diameter, with a rather sharp 
odor of trim ethyl amine; calyx more or less lanate, later glabrescent, its 
teeth with glandular cilia on the margin; petals white, orbicular, 4 — 5 mm 
long, the upper surface lanate, pubescent from base; stamens 20, as long 
as petals; styles 3 (less often 2, 4, or 5), free, hairy in lower part; fruits 



377 



289 



378 



subglobose, bright red at maturity, 9 — 10 mm in diameter; seeds usually 3, 
narrowly oblong, acute, reddish. Fl. May— July, fr. September. Forest 
edges, glades, undergrowth, less often meadows, also rocky or stony sites, 
riverbanks bluffs, etc.; often planted near houses and in orchards.— 
European part: Kar. -Lap., Dv. -Pech. (W. part), Lad. -Ilm., U. V., V. -Kama, 
U. Dnp., M. Dnp., V. -Don, Transv., Crim.; Caucasus: Cisc, Dag., E. and 
S. Transc, Tal. Gen. distr.: . Arc. (Iceland), Scand., Centr. Eur.; Atl. Eur., 
Bal. -As. Min. Described from N. Europe. Type in London. 

Economic importance. Wood fine-grained, reddish, lustrous, hard, 
polishes well, furnishes good material for turning and furniture, etc., 
mainly for machine parts such as blocks, rollers, and cogs, requiring great 
strength. The bark contains 7.26% tannin compounds (Wehmer I, 448), the 
fruits malic and citric acids, 4 — 8% dextrose, also traces of hydrocyanic 
acids, (harmless); used for infusions, vinegar, vodka, marmalade, jam, 
pastilles, etc. The seeds contain 21.9% fatty oils and the glucoside 
amygdalin; the young branches furnish a black dye. Reproduces in 
cultivation by roots suckers. 

Note. The Caucasian Sorbus with smooth, rather firm leaves forms 
"islands" among S. aucuparia, and will be separated as an independent 
species if distinguishing characters can be ascertained. 

5. S. glabrata Hedlund in Sv. Vet. Akad. Handl. 3 5, I (1901) 45.— 

S. aucuparia var. glabrata (Hedl.) Perfil'ev, Fl. Sev. Kr. II, 173 (non 
Wimm. et Grab.). 

Shrub 1— 2 m high; winter buds densely sericeous-pubescent; leaflets 
acute, tapering to a sharp point, firm, with very sparse scattered pubescence 
on both sides, acutely serrate; common petiole, petiolules and angles 
of veins much less pubescent below than in S. aucuparia, but peduncle 
often densely lanate; inflorescence 6 — 8 cm in diameter, with strongly 
declinate branches; flowers often large, to 12 mm in diameter; calyx teeth 
longer than broad, with rounded margins, fruits slightly elongated. Fl. July, 
fr. September. 

River valley terraces and slopes. — Arctic: Arc. Eur. Gen. distr.: 
N. Scand. As far east as the Bol'shezemel'skaya Tundra (Andreev). 
Described from the Kola Peninsula. Type in Lund. 

Note. In his monograph (p. 45), Hedlund confuses this tree with the 
smooth-leaved form S. aucuparia L. from Central Europe, first recorded 
by Gilib. in his Fl. lithuan. V, 233. 

6. S. polaris Koehne in Fedde, Repert. X (1912) 502. 

Shrub 1 m or more high; branches ca.3mm in diameter; buds ca.3mm 
long, often densely white -tomentose in upper part; stipules small, lanceolate, 
caducous; leaves together with petiole 1.5 — 2 cm broad, 7 — 12 cm long, 5- or 
6 -paired; common petiole wingless, slender, with long hairs on upper surface 
at base of leaflets, otherwise glabrous or subglabrous, the intervals 
between leaflets 0.8 — 1.4 cm long; leaflets diminishing upward and downward, 
broadly lanceolate, the middle leaflets 3— 4cm long, 0.7 — 1.3 cm broad, 
acute, entire from base, acutely serrulate higher up, with 8 — 20 teeth on each 
side, paler below, green, smooth, with 12 — 16 slender, inconspicuous veins 
on each side; inflorescence 2.5 — 3 cm in diameter, 4.5 — 5 cm long, glabrous 
or scattered-hairy, few-flowered; calyx glabrous or subglabrous, its teeth 



290 




PLATE XXIII. 1— Sorbus amurensis Koehne: a) flower, b) fruit, c) stipule, d) bud; 
2-S.schneideriana Koehne: a) flower, b) fruit, c) stipule, d) bud. 



291 



381 



with few hairs; petals 3 mm long, broadly rounded, with beards above from 
base, as long as stamens; carpels 3, rarely 4, with hairy apex. Fl. end of 
June — July, fr. September. (Plate XXIV, Figure l). 

River valley terraces and slopes.— Arctic: Arc. Sib. Described from 
the Shchuch'ya River, north of the Ob River mouth and the Arctic Circle. 
Type in Germany. In the Leningrad Herbarium these are closely related 
forms from the Polui and Nadnya rivers, which — like the Shchuch'ya River — 
flow into the Ob Gulf but farther east. 

Note. The Arctic form is easily distinguished from S. sibirica Hedl. 
by its small inflorescences, impoverished pubescence, short stamens, and 
number of carpels. 

7. S. sibirica Hedl. in Sv. Vet. Akad. Handl. 35, I (1901) 44; Perfil'ev, Fl. 
Sev. Kraya. II 173; Koehne in Fedde, Repert. X (l 912) 502. — S. au cup aria 
ssp. sibirica (Hedl.) Kryl., Fl. Zap. Sib. (1933) 1464.— Pyrus aucu- 
paria Turcz., Fl. baic. -dah. 11,412. 

Tree 3 — 10 m high; winter buds glabrous or covered with short, scattered, 

slender hairs; summer buds and the young branches pubescent; leaves 

10 — 20 cm long, 8 — 12 cm broad; leaflets oblong -lanceolate, 3.5 — 5 cm long, 

1—1.5 cm broad, green and smooth above, gray-green with papillate epiderm 

below, more or less pubescent along the midrib, with 5 — 10 pairs of lateral 

veins, serrate-dentate; inflorescences dense, broad, 6 — 8 cm long, 8 — 12 cm 

i, & 
broad, their branches glabrous or scattered-hairy; stamens /4 longer 

than white orbicular petals; corolla 7 — 9 mm in diameter; calyx subglabrous, 

with broadly deltoid teeth; pistils 3 or 4; apex of ovary and base of style 

long-hairy. Fl. June, fr. August-September. 

Forest- tundra, forest zone to edge of steppe, coniferous and deciduous 
mountain forests. — Arctic: Arc. Eur. and Sib. European part: Dv. -Pech., 
V. -Kama; W. Siberia: Ob, U. Tob. (NE part), Irt., Alt.; E. Siberia: Yenis., 
Lena-Kol., Ang. -Say., Dau.; Far East: Ze. -Bu. and Uss. Gen. distr. : Mong. 
(N. mountains), the northern limit, according to Krylov, on the Yenisei 
69°40' N, in the Urals 61°45'N., the southern limit 48°. Described from 
Siberia, probably from Asinovo village on the Yenisei. 

Note. A species replacing S. aucuparia L. but well differentiated 
from its European forms. The comparison with S. glabra Gilib. is an 
unfortunate one. Sorubus al t ai c a, described by E. Koehne (in Fedde, 
Repert. X, 1912, 505) after the very unsufficient material collected by 
O. Duhmberg in 1881 in Barnaul, differs in the shape and serration of its 
leaflets, the absence of papillae on their lower surface, and the presence of 
3 carpels. No one else reports this form, and there is no reason to think 
that there is a separate species of mountain ash growing near Barnaul. 
The same can be said about S. au cu p a r i a var. du mb e r g i i Koehne in 
Fedde, Repert. X (1912) 504. Both these plants have been defined by 
C. K. Schneider as S. sibirica. The originals — Duhmberg Nos. 996 and 
997 — are preserved in Berlin. 

8. S. amurensis Koehne in Fedde, Repert. X (1912) 513; Nakai in Fl. 
Sylv. Koreana VI (1916) 22. —Pyrus aucuparia Maxim., Prim. Fl. Am. 
(1859) 103. — Sor bus aucuparia Kom., Fl. Manshur. II, 472. — Ic. : 
Nakai, I.e., tab. 2. 



292 



Tree 4 — 8 m high; bark gray with dark, numerous, horizontal lenticels; 
young branches reddish with white linear or oblong lenticels, 2.5 — 5mm 
in diameter; buds more densely sericeous -lanate in their upper part some 
stipules linear, caducous, others broad, coarsely dentate, green; leaves 
together with petiole 2.8 — 3.5cm broad, 15.5 — 21 cm long, 5 — 7-paired; 
common petiole slender, wingless, hairy in places, but smooth in larger part; 
lower leaflets petiolulate, the upper sessile, linear or broadly lanceolate, 
4 — 5.5 cm long, 1.3 — 1.6 cm broad, entire from base (some to the middle), 
„ R9 acutely serrulate in upper part, with a long or short sharp point, smooth and 
dark green above, pale gray-green and glabrous or slightly hairy at base 
below, the lateral veins 15 — 20 pairs, inconspicuous; inflorescence ca 13 cm 
in diameter, 11 cm long, glabrous or slightly hairy; calyx glabrous or hairy 
with ciliate teeth; petals 4 mm long, orbicular, lanate-barbate at base 
above, slightly shorter than stamens; styles 3, lanate at base; fruits bright 
red, globose, 6 — 7 mm in diameter. Fl. May — June, fr. September — October. 
(Plate XXIII, Figure l). 

Mountain forests, especially on rocky slopes and among large stones, 
along streams and on afforested river islands.— Far East: Ze. -Bu., Uda, 
Uss. Gen. distr. : Jap. -Ch. (Korea-Manchuria). Described from the Amur 
after Maximowicz's specimen. Type in Berlin; cotype in Leningrad. 

Note. In the work by Kom. and Alis., Opred. rast. Dal'nevost. Kraya II, 
637, I have recorded Sorbus discolor Maxim, for the Ussuri region, 
with the indication that its leaves are not acutely serrate, but have only a 
few teeth near the apex; I have added that they are firmer and more obtuse 
than the leaflets of S. amurensis. The diagnosis of Maximowicz 
(Primitiae I.e., note to p. 103) is very incomplete; later Rehder (Manual 
Cult. Trees and Shr. N. Amer., 1927, 379) gave a more detailed description 
of this species, adding as a synonym also S. pekinensis Koehne, 1906, 
in Mitt. Deutsch. Dendr. Ges.; it appears, however, that this species has 
white flowers. In view of this, and also because Nakai indicates only 
S. amurensis for Korea, I think now that the S. Ussuri ash belongs to 
S. amurensis Koehne. 

9. S.boissieri C. K. Schn. in Bull. Herb. Boiss. VI, 1906, 312; Ej. Ill, 
Laubholzk. I, 671 ; Grossg., Fl. Kavk. VI, 287. 

Tall shrub; branches more or less pubescent, the annotinous branches 
brown-red, glabrous or with remains of hairs; lenticels thin, light; buds 
fusiform, glabrous, more or less glutinous;, leaves 5 — 7-paired, to 30 cm 
long, with scattered hairs and pubescence at base of leaflets along the midrib; 
leaflets dark green above, whitish-glaucous below, dentate only above the 
middle, short-acuminate, 5 — 7 cm long, 1.8 — 2.5 cm broad; inflorescence 
many-flowered, to 15 cm in diameter, glabrous or subglabrous; flowers 
10 — 12 mm in diameter; stamens 20; petals broadly ovate; styles 4 or 5; 
fruits ovoid-rounded, 8 — 9 mm broad, yellowish red. Fl. May, fr. September. 

Upper timberline, mountains. — Caucasus: W. Transc Gen. distr.: As. 
Min. Described from Lazistan; reported also for Abkhazia and Adzharia. 



383 



10. S. commixta Hedl. in Sv. Vet. Akad. Handl. 35, I (1901) 38; C. K. Schn., 
III. Laubholzk. I, 677. — S. c ommixt a var. Takasui Kudo in Journ. 
Coll. Agr. Univ. Hokkaido XII (1923) 42.— Pyrus aucuparia Fr. Schmidt 
in Bull. Ac. Sc. Petersb. V (1862) 140. 



293 



384 



Shrub or tree 4 — 8 m high; bark dark gray, lenticels oblong or subulate, 
horizontal; branches quite smooth, gray, annotinous branches red-brown; 
buds glabrous, sticky, 1.6 cm long, acute; leaves smooth, 4 — 6 -paired; 
leaflets lanceolate or linear-lanceolate, mucronate, deeply serrate or 
biserrate, often with aristate teeth, paler below than above; inflorescence 
quite devoid of pubescence, dense; flowers ca 1 cm in diameter; fruits 
bright red, 5 — 7 mm in diameter; calyx teeth (corona) inflexed. Fl. June, 
fr. August. 

Mountain forests.— Far East: Sakh.; Gen.distr.: Jap. -Ch. Described 
from Japan. Type in Stockholm or Helsinki. 

11. S. kamtschatcensis. Kom. in Notulae systematicae ex Herb. Horti Bot. 
Petrop. II (1921) 135; Fl. Kamtsch. II, 239; E. Hulten, Fl. of Kamtch. II, 45. 

Tree 2 — 12 m high, smooth, with dark gray bark; branches somewhat 
thickened; leaves dull, dark green above, pale green below, 10 — 24 cm long; 
petioles slightly pubescent at base and at point of insertion, often red; 
leaflets (mostly) 6 pairs conspicuously petiolulate, lanceolate or oval- 
lanceolate, mucronate, acutely serrate (serration always simple), erect, 
short, 4 — 9 cm long, 10 — 20 mm broad, the lateral veins 15 — 18 pairs, 
depressed above; inflorescence ca 9 cm in diameter; bracts falcate, short; 
flowers white, aromatic, ca. 1 cm in diameter; sepals erect, quite glabrous; 
fruits small, oblong, light red, bitter; petals oblong, rounded at the apex, 
ca. 4 mm long; stamens as long as petals; styles mostly 3. Fl. June, 
fr. September. 

Common birch forests, mostly along river valleys: solitary.— Far East: 
Kamch. Endemic. Described from the left bank of the Avacha River below 
Elizov; fruits from the Levaya Shchapinskaya River. Type in Leningrad. 

12. S. a§adyrensis Kom. in Not. syst. ex Herb. Horti Bot. Petrop. II (1921) 
134. 

Erect shurb with smooth bark and slender branches; buds narrow, acute, 
with long soft hairs on scale margins; leaves 7 — 18 cm long, quite glabrous 
or with few hairs on the petioles; leaflets usually 6 -paired lanceolate with 
cuneate base, paler below, 1.5 — 5.5 cm long, 0.6 — 1.7 cm broad, dull above, 
with inconspicuous network of veins, slightly coriaceous, serrate; 
inflorescence compact, 3— 4cm in diameter, glabrous; bracts linear; 
pedicels ca. 3 mm long; calyx glabrous, with straight eciliate teeth shorter 
than calyx; petals orbicular, 2.5 — 3 mm long; stamens very short; fruits 
light red, globose, 5 — 8 mm in diameter; seeds elongated, slightly curved? 
Fl. July, fr. September. 

Arctic: An. Endemic. Described from the middle reaches of the Anadyr 
River (mouth of its tributaries the Snezhnaya and Main). Type in Leningrad. 



Series 3. Tianschanicae Kom.— Leaflets green below; winter buds more 
or less pubescent; leaves typically 6 -paired; flowers and fruits larger than 
in species of the series Aucupariae. 

13. S. tianschanica Ruprecht in Mem. acad. Petersb., ser. 7, vol. XIV 
(1869) 46; C. K. Schneider, III. Laubholz. I, 668; Fedtsch., Consp. Fl. Turk, 
in Beih. Bot. Centr. XXIV (1908) 198.— Pyrus tianschanica Franchet 
in Ann. Sc.nat. XVI (1883) 267. 



294 




PLATE XXIV. 1-Sorbus p ol ar i s Koehne: a) fruit. -b) stipule, c) bud; 2-S.t i ans ch a ni a 
Rupr.: a) fruit, b) stipule, c) bud. 



295 



387 



Tree 3 — 5m high or shrub; young branches olive-brown or red-brown, 
slightly pubescent, the lenticels inconspicuous; buds acute, densely 
pubescent, sometimes subglabrous; leaves typically 6 -paired, 10 — 14(22) 
cm long, quite glabrous or with hairy petioles; leaflets slightly coriaceous, 
lanceolate, mucronate, with 16 — 18 erect, appressed, short teeth on each 
side, light green below; inflorescence to 15 cm in diameter, with red 
glabrous branches; flowers 1.5 — 2 cm in diameter; stamens short; ovary 
densely lanate at the apex; styles mostly 5, sometimes 2 — 4; mature 
fruits dark red with glaucous bloom, to 12 mm in diameter. Fl. May — July, 
fr. August -September. (Plate XXIV, Figure 2). 

Mountain forests and shrub thickets near the upper timberline. — Centr. 
Asia: T. Sh., Pam. -Al., Dzu. -Tarb. Gen. distr. : Dzu. -Kash. Described 
from the Shamsi and Mulda-ashu gorges between the Tokmak River and 
the upper reaches of the Narym River (district of Lake Son-kul). Type 
in Leningrad. 

Notel. S. tianschanica is sharply distinguished from all 
i-ri>rc,scnl..-iL.iv<\s of the; series A u c u p a r i a e, and theref ore I deem it 
necessary to acknowledge it as the representative of a separate series of 
species. For the time being it remains isolated, but a general monographic 
treatment of the genus So r bus should also include some additional 
southern species. 

Note 2. In Hedlund's monograph (Sv. Vet. Akad. Hadling, 35, I, 39 — 40) 
two additional American species are recorded for Kamchatka: S. pumila 
Rafin. and R. sitchensis Roemer, which Soviet researchers have not 
yet been able to discover in Kamchatka. Their occurrence in Kamchatka 
is most unlikely, and the author's error has probably been caused by a 
confusion of labels. 



Subgenus 2. Hahnia Medik., * Gesch. d. Bot. (1793) 81, pro gen. — Leaves 
simple, entire or dentate- lobate or lobate, only the lowest pair 
of lobes sometimes quite separate from the others. Styles 2 — 5, nearly 
always connate at base. Carpels in middle part mostly completely connate, 
variable in degree of union with the hypanthium; ovary sometimes apparently 
wholly inferior. Cell walls hard. 

1. Leaves pinnately compound or pinnatisect, with 1 or 2 pairs of leaflets 
or segments, simple at the apex 2. 

+ Leaves simple 4. 

2. Leaves pinnatisect, with 1 or 2 pairs of segments, white-tomentose 
below 32. S. turkestanica Hedl. 

+ Leaves pinnatisect (nearly pinnately compound at base), with 2 or 3 

pairs of segments, slightly tomentose below, gray-green 3. 

3. Leaves gray-green above 33. S. dualis Zinserl. 

+ Leaves green above *S. fennica Fr. 

4. Leaves green on both sides, glabrous below, with 3 — 5 more or less 

acute lobes, sometimes pinnatipartite below 

34. S. torminalis Crantz. 



I'iv, II iik ill |, v Vn.l >.. 1 1 1 ; . ■ 1 liny,. 



296 



+ Leaves more or less strongly pubescent below, if glabrous then 

unlobed " 

5. Leaves greenbelow, glabrous or slightly pubescent (not tomentose); 
calyx caducous in mature fruits 

+ Leaves tomentose below 

6 Leaves glabrous below or slightly pubescent along the veins .... 

14. S. subfusca (Ldb. ) Boiss. 

+ Leaves slightly pubescent below (not only along the veins) 

15. S. albovii Zinserl. 

Q 

7. Leaves distinctly lobed 

+ Leaves unlobed or obscurely lobed 12. 

8. Leaf lobes shallow (not more than % the half-width of blade); 

lateral veins 5 — 7 pairs; leaves white-tomentose below, with 

obliquely-triangular teeth, the midrib and lateral veins forming an 

angle of 45° 31. S. persica Hedl. 

9 
+ Lateral veins 7-9 pairs 

9. Leaf lobes with conspicuous teeth on both margins 10- 

+ Leaf lobes entire along inner margin or with 1 or 2 small teeth only 

at the apex 

Angle formed by midrib and lateral veins usually more than 45°; 
teeth of lobes and those in upper, unlobed part more or less equal 

23. S. armeniaca Hedl. 

+ Angle formed by midrib and lateral veins usually less than 45°; 

teeth in upper unlobed part of leaf much larger than those of lobes 

*S. scandica Fr. 

11. Leaves ovate-rounded or broadly elliptic (l .2 - 1 .3 times longer 

than broad) 21. S. caucasica Zinserl. 

+ Leaves elliptic or obovate-elliptic (1.4-1.7 times longer than broad) 
22. S. woronowii Zinserl. 

12. Leaves elliptic, ovate-elliptic, or slightly obovate-elliptic (1.5 — 2 times 
longer than broad), rounded or slightly narrowed at base, dingy gray-green 
below, slightly tomentose, distinctly duplicato-dentate with subobtuse 
teeth of the first order; calyx caducous in mature fruits 

17. S. colchica Zinserl. 

+ Leaves white or gray-tomentose below or, if grayish green and 

slightly tomentose, then not more than 1.5 times as long as broad and 
teeth of the first order not subobtuse 13 - 

13. Leaves grayish green or gray below, those on sterile shoots narrow, 
fusiform-elliptic, fusiform-lanceolate, or elliptic -lanceolate, with 
cuneate base, sharply differentiated from other leaves, which are 
broader and without cuneate base 14 - 

+ Leaves white or gray below, those on sterile shoots not sharply 

differentiated from other leaves (if differentiated then leaves white 
below) 15 ' 

14. Leaves very coriaceous, with 10 — 11 pairs of veins, strongly 
impressed above, 1.2 — 1.3 times as long as broad; buds ca. 10 mm 
long 19. S. buschiana Zinserl. 

+ Leaves softer, with 7-9 pairs of veins, slightly impressed above, 

1.4 — 1.5 times as long as broad; buds 3 — 5 mm long 

20. S. schemachensis Zinserl. 



297 



389 



15. Leaves grayish green, rounded-elliptic, less often obovate-elliptic, 

slightly tomentose; calyx caducous in mature fruits 

16. S. subtomentosa (Alb.. ) Zinserl. 

+ Leaves white or gray, more or less strongly tomentose 16. 

16. Leaf teeth (simple or of the second order) numerous, not less than 

20 on each side 19. 

+ Leaf teeth less numerous (not more than 1 5 on each side), usually 

larger 17. 

17. Teeth more or less obtuse (sometimes passing into crenae); leaves 
obovate 30. S. obtusidentata Zinserl. 

+ Teeth acute 18. 

18. Leaves orbicular or broadly ovate or obovate-elliptic, rounded 
toward base, usually rounded at the apex 2 8. S. turcica Zinserl. 

+ Leaves obovate, obovate-elliptic, or subrhomboid, more or less 

acuminate, conspicuously tapering toward base 

2 9. S. taurica Zinserl. 

19. Leaves orbicular, rounded-elliptic, obovate, or obovate-elliptic, 
1—1.4 times as long as broad, white -tomentose below, with acute 
teeth 23. 

+ Leaves gray-tomentose below, if white-tomentose then lateral veins 

6 — 8 pairs 20. 

20. Leaves strongly pubescent, with strongly impressed veins above; 

leaf teeth subobtuse; calyx caducous in mature fruits 

18. S. velutina (Alb. ) C. K. Schn. 

+ Leaves without strongly impressed veins; teeth acute; calyx 

persistent in mature fruits 21. 

21. Leaves narrow (about twice as long as broad), small, with 6 — 8 pairs 
of veins, white-tomentose below 27. S. baldaccii Deg. et Fritsch. 

+ Leaves broader, large, with 8 — 12 pairs of veins 22. 

22. Leaves cuneately tapering toward base, with 8 — 10 pairs of veins 
24. S. kusnetzovii Zinserl. 

+ Leaves not cuneately tapering toward base, with 10 — 12 pairs of 

veins *S. aria Crantz. 

23. Leaves larger, 5 — 10 cm long, with white-tomentose veins below. 
Tree (or large shrub?) 25. S. graeca (Spach) Hedi. 

+ Shrub 50 — 80 cm high; leaves small, 4 — 5 cm long, the veins slightly 

pubescent, very prominent on the whitish blade 

26. S. migarica Zinserl. 



Section 1. ARIA DC, Prodr. II (1825) 635.— Flesh of fruit without grit 
cells; carpels free at the apex. 



Series 1. Subfuscae Zinserl.— Sepals caducous in mature fruits. Leaves 
dentate or duplicato-dentate, green below, subglabrous or slightly pubescent 
or else tomentose and gray-green or gray; teeth 30 or more on each side; 
lateral veins 8 — 11 pairs (in S. schemachensis 7 — 9 pairs) forming 
a 45° angle with the midrib. 



298 



14. S. subfusca (Ldb.) Boiss., Fl. Or. II (1872) 659. - Hedlund, Monogr. 
(1901) 109; C. K. Schn., Handb. d. Laubholzk. J, 685.— Crataegus 
subfusca Ldb. apud Nordm., Vorl. Diagn. in Bull, de l'Acad. de St. Petersb. 
II (1837) 313 et Fl. Ross. II (1844) 87.— Aria Szovitsii Decaisne, Mem. 
Pom. in Nouv. arch, du mus. l'hist. nat. Paris X (1874) 165.-Sorbus aria 
var. glabra Albow, Prodr. fl. colch. (l 895) 71 .— Ic. : C K. Schn., 1. c. 

Tree; buds subglabrous (scarcely pubescent on the margin); leaves 
obovate- elliptic to broad-elliptic, tapering or rounded toward base, acute 
or subobtuse, mucronate, 6.5 — 11 cm long, 3.8 — 7 cm broad (length- to- width 
ratio 1.3 — 2.2), with 8 — 11 pairs of veins slightly impressed above; the 
blade glabrous on both sides (sometimes slightly pubescent only along 
midrib), duplicato-dentate, less often almost simply dentate, with acute teeth 
almost to base of blade; pedicels glabrous or subglabrous at anthesis; calyx 
glabrous, with acute linear-lanceolate teeth, more or less erect at anthesis, 
later declinate, slightly tomentose on the margin; petals oblong-ovate or 
rounded -ovate, 1.5 — 2 times as long as calyx; styles 2, approximate; 
fruits ovoid or globose, red, later turning blue. Fl. June — July, fr. August- 
September. (Plate XXV, Figure 6). 

Mountains, atthe upper timberline, at 1,700 — 2,100 m. — Caucasus: Cisc. 
(Teberda Reserve), W. Transc. (Adzharia, Guria, Mingrelia, Svanetia), 
E. Transc. (S. Ossetia). Endemic. Described from the Caucasus. Type 
in Leningrad. 

15. S. albovii Zinserl. sp.n. in Addenda VIII, p. 379. — S. aria var. 
Concolor Albow, Prodr. fl. colch. (1895) 71.— S. a r i a var. concolor 
Boiss., Fl. Or. II (1872) 658, p. p. — S. c one ol o r C K. Schn., Handb. d. 
Laubholzk. I (1906) 686 p. p.; Grossg., FL Kavk. IV (1934) 288.— 

Tree; buds glabrous or slightly tomentose; leaves obovate -elliptic or 
elliptic (rarely rounded-elliptic), somewhat narrowed, less often rounded at 
base, subacute or obtuse, usually mucronate, 7 — 10 cm long, 4 — 7 cm broad 
(length- to- width ratio 1.2- 1.6), with 8-11 pairs of lateral veins, glabrous 
above (or when young slightly pubescent along the veins), green and very 
slightly pubescent below (sometimes only along midrib and lateral veins), 
dentate, usually obscurely duplicato-dentate in upper part, with acute teeth, 
nearly to base of blade; pedicels glabrous or slightly tomentose; calyx 
tomentose, with triangular-lanceolate acute teeth; petals rounded -ovate; 
styles 2; fruits globose-ovoid, red, turning blue. Fl. July, fr. August. 

Upper parts of the forest zone (beech and birch forests) and in the 
subalpine zone at 1,800 — 2,000 m. — Caucasus: Cisc. (w. part) W. Transc. 
(Abkhazia), S. Transc. (Zangezur), E. Transc. (S. Ossetia). Endemic. 
Described from the Caucasus. Type in Leningrad. 

16. S. subtomentosa (Alb. ) Zinserl. sp. n. in Addenda VIII, p. 379. — 
S. aria var. s ub t om en t o s a Albow, Prodr. fl. colch. (l 895) 71; Vol'f 

i Palibin, Opred. der. i kust. (l 904) 462. — S. c one o 1 o r C. K. Schn. Handb. 
d. Laubholzk. I (1906) 686 p. p. 

Tree; buds tomentose on the margin, less often glabrous; leaves 
rounded-elliptic, less often obovate- elliptic, rounded or slightly narrowed 
at base (less often strongly narrowed), obtuse or subacute, usually mucronate, 
6 — 13.5 cm long, 4.5 — 10.5 cm long (length-to-width ratio 1.1—1.5), with 
8 — 11 pairs of lateral veins slightly impressed above, the blade glabrous 
(or slightly pubescent only along midrib) above, slightly tomentose below, 



299 



392 



grayish green, dentate, duplicato-dentate in upper part, with acute teeth 
nearly to base of blade; pedicels glabrous or slightly tomentose; fruits 
globose-ellipsoid, reddening; calyx with triangular-lanceolate, spreading, 
tomentose teeth in immature fruits. Fl. unknown, fr. August. (Plate 
XXV, Figure 7). 

Subalpine zone, at upper timberline, sometimes beech forests, at 
1,900 — 2,200m. Caucasus: Cisc. (mainly in the west, as far east as 
Chirkei), W. and S. Transc. (Akhalkalaki). Endemic. Described from the 
Caucasus. Type in Leningrad. 

17. S. colchica Zinserl. sp. in Addenda VIII, p. 380. 

Tree; buds slightly tomentose on the margin; leaves elliptic, ovate- 
elliptic, or slightly obovate-elliptic, rounded or slightly narrowed at base, 
obtuse or subacute, usually mucronate, 8.5 — i cm long, 5 — 7 cm broad 
(length- to- width ratio 1.5 — 2), with 10— 11 pairs of lateral veins, the blade 
glabrous (or slightly pubescent along veins) above, dingy gray-green and 
slightly tomentose below, conspicuously duplicato-dentate in upper j 2 or 
% of blade, dentate lower down, nearly to base; leaf teeth of the first order 
subobtuse (apiculate), with long lower and short upper margins, teeth of 
the second order acute; pedicels glabrous; fruits globose, red, later 
turning blue. Fl. July, fr. September. (Plate XXV, Figure 4). 

At the upper timberline. — Caucasus: W. Transc. (Circassia, Abkhazia, 
Adzharia). Endemic. Described from the Caucasus. Type in Leningrad. 

18. S. velutina (Alb.) C. K. Schn., Handb. d. Laubholzk. I (1906) 686, 
Grossg., Fl.Kavk. IV (1934) 288 p.p. —S. aria var. velutina Albov, Prodr. 
fl. colch. (1895) 71.— Ic: C. K. Schn., 1. c, I (1906) 686, f. 377 d. 

Tree; buds tomentose -pubescent; leaves coriaceous, obovate-elliptic, 
rounded or slightly narrowed at base, obtuse or subacute, sometimes 
mucronate, 6 — 12 cm long, 5 — 9 cm broad (length to width ratio 1.2 — 1.4), 
with 9 — 11 pairs of lateral veins strongly impressed above, the blade 
glabrous or slightly pubescent along the veins above, densely gray-tomentose 
below, dentate in lower part and duplicato-dentate at the apex, subentire in 
lower third leaf; teeth subobtuse; pedicels tomentose; calyx tomentose 
with spreading (in immature fruits) triangular-lanceolate teeth; flowers 
unknown; fruits globose-ellipsoid; red, later turning blue. Fr. October 
(Plate XXV, Figure 10). 

Subalpine zone: in shrub thickets and in meadows at 1,750 — 1,900 m. 
Caucasus: W. Transc. (Circassia and Abkhazia). Endemic. Described 
from the Caucasus. Type in Leningrad. 

19. S. bushiana Zinserl. sp. n. in Addenda VIII, p. 381. 

Tree; buds slightly tomentose, large, to 1 cm long; leaves coriaceous, 
glabrous above (except midrib), gray-green-tomentose below, broadly 
elliptic or ovate -elliptic, rounded at base, obtuse or subacute, 10 — 11 cm 
long, ca. 8.5 cm broad (length-to-width ratio 1.2 — 1 .3), with 10 — 11 pairs of 
lateral veins, the blade obscurely duplicato-dentate nearly to base, with 
acute teeth; some leaves on sterile shoots narrow, fusiform and narrowly 
elliptic or fusiform-lanceolate, acuminate, with cuneate base; pedicels 
subglabrous; calyx teeth triangular-lanceolate, white-tomentose; flowers 
and mature fruit unknown. Immature fruits in August. (Plate XXVI, 
Figure 9). 



300 




PLATE XXV. 1-Sorbus caucasica Zinserl., leaf; 2— S.persica Hedl., leaf; 3-S.ar- 
meniaca Hedl., leaf; 4-S.colchica Zinserl., leaf; 5-S.turkestanica Hedl.; 6-S.sub- 
fusca (Ldb.)BoiSS.,leaf:a) fruit; 7 -S.sub foment os a (Alb.) Zinserl. .leaf; 8-S.scandica 
Fr., leaf margin; 9-S.wo ronowi i Zinserl., leaf; 10 -S.velutina (Alb.) C. K. Schn.,leaf. 



301 



Rock streams at 1,850 — 2,200 m. — Caucasus: E. Transc. (S. Ossetia, 
near the village of Nizhnii Ermani). Endemic. Described from the 
Caucasus. Type in Leningrad. 

20. S. schemachensis Zinserl. sp. n. in Addenda VIII, p. 381. 

Tree; buds slightly tomentose -pubescent, 0.3 — 0.5 cm long; leaves 
coriaceous, glabrous above (except midrib), gray-tomentose below, elliptic, 
obovate-elliptic, or ovate-elliptic, rounded at base, obtuse, 7 — 8.5 cm long, 
5.5 — 6 cm broad (length-to-width ratio 1.4 — 1 .5), with 7 — 9 pairs of lateral 
veins, the blade dentate nearly to base (double dentation scarcely discernible) 
with small, acute teeth; some leaves on sterile shoots much narrower, 
elliptic -lanceolate, cuneately tapering toward base, sometimes acuminate; 
pedicels white-tomentose. Flowers and mature fruits unknown; immature 
fruits globose-ovoid. Immature fruits in August. (Plate XXVI, Figure l). 

Limestone rocks at ca. 1,150 m. — Caucasus: E. Transc. (Myudusa in 
the Shemakha District). Endemic. Described from the Caucasus. Type 
in Leningrad. 

Note. Closely related toS. buschiana in the sharply differentiated 
double shape of leaf; possibly the result of a cross between S. busch- 
iana and S.graeca. 



Series 2. Lobatae Zinserl.— Sepals persistent in mature fruits. Leaves 
conspicuously lobate, gray-tomentose below; teeth 30 — 40 or more on each 
side; lateral veins 7 — 9 (10) pairs, forming an angle of ca. 45° with the 
midrib. 

21. S. caucasica Zinserl. in Not. syst. herb. H. B. P. IV (1923) 17-18; 
Grossg., Fl. Kavk. IV (1934) 283.— S. scandica Lipsky, Fl. Kavk. I, 296 and 
Suppl. 51 et al. auct. fl. cauc norm, (non Fr.). 

Tree; buds pubescent; leaves ovate-rounded or broadly elliptic, more 
or less conspicuously tapering toward base, obtuse, less often short- 
acuminate, 8 — 13 cm long, 6.5 — 11 cm broad (length-to- width ratio 1 .2—1 .3), 
gray-tomentose below, pubescent only along midrib above, 5 — 7-lobed 
(lobes passing into teeth at the apex), with 7 — 9 pairs of lateral veins forming 
an angle of ca. 45° with the midrib; leaf lobes oblong, subobtuse, apex and 
in lower part dentate, in upper part entire, less often some lobes with 
1 or 2 small teeth in upper part, the lowest and the following lobes to 
half the half-width of blade; leaf teeth 30 — 50 on each margin, acute, those 
in upper part of leaf much larger than the others; pedicels tomentose; 
calyx teeth acute, triangular, tomentose, erect in fruit; petals obovate; 
styles 2 or 3; fruits ovoid or globose, red, later turning blue. Fl. May- 
June, fr. August. (Plate XXV, Figure l). 

Upper part of forest zone, mainly on rocks, at 900 — 1,600 m. — Caucasus: 
Cisc, Dag., E. and S. Transc. (rarely). Endemic. Described from the 
Caucasus. Type in Leningrad. 

22. S. woronowii Zinserl. sp. n. in Addenda VIII p. 381.— ? S. aria var. 
incisa Albov, Prodr. fl. colch. (1895) 72. 

Tree; buds pubescent; leaves elliptic or obovate-elliptic, slightly 
narrowed toward base, short-acuminate, 9 — 12cm long, 7 — 8.5cm broad 



302 



397 



(length-to-width ratio 1.4 — 1.7), 4- or 5-lobed (lobes passing into teeth at 
the apex), pubescent only along midrib above, gray-tomentose below, with 
7 — 9 pairs of lateral veins forming an angle of ca. 45° with the midrib; 
leaf lobes oblong, acuminate, in lower part dentate, in upper part entire or 
in some lobes sometimes with 1 or 2 small teeth, the two lower pairs of 
lobes to lz of half-width of blade; leaf teeth 30 — 40 on each margin, acute, 
much larger at the apex than lower down; pedicels tomentose; calyx teeth 
acute, triangular, tomentose, not spreading in fruit; styles 2; fruits ovoid, 
red, later turning blue. Fl. unknown, fr. September. (Plate XXV, Figure 10). 

Limestone rocks. — Caucasus: W. Transc. (Mt. Fisht). Endemic. 
Described from the Caucasus. Type in Leningrad. 

*S. scandica Fr., Fl. Holland, (l 81 7) 38; Grossg., Fl. Kavk. IV (l 934) 289. 

Tree, very similar to the preceding species, from which it is distinguished 
by leaf lobes with always 1 or 2 teeth in upper part similar in size to those 
in lower part (see Plate XXV). Not growing wild in the USSR (distributed in 
countries bordering on the Baltic Sea), sometimes cultivated in parks. 

23. S. armenica Hedl., Monogr., d. Gatt. Sorbus (1901) 69; C. K. Schn., 
Handb.d. Laubholzk. I (1906) 693.— Ic: Hedl., Monogr. d. Gatt. Sorbus, 69, 
Fig. 17. 

Tree; buds pubescent; leaves ovate-elliptic, elliptic, or oblong -elliptic, 
rounded toward base, acute or subobtuse, 6.5 — 7.5 cm long, 4 — 5.5 cm broad 
(length-to-width ratio 1.2 —1.8), pubescent only along midrib above, gray- 
tomentose below, with 7 — 9 (10) pairs of lateral veins forming at 45° angle 
with midrib, 5- or 6-lobed (lobes passing into teeth at the apex); leaf lobes 
considerably broadened (rounded-triangular) toward base, dentate on 
both margins, the two lowest pairs V3 — % half-width of of blade; leaf teeth 
30 — 50 on each margin, acute, more or less similar in upper (unlobed) part 
of blade and lower; pedicels tomentose; calyx with acute triangular teeth, 
tomentose; fruits ovoid (mature fruits unknown). Fl. June. (Plate XXV, Figure 3). 

Upper timberline, at ca. 1, 500 m. — Caucasus: S. and E. Transc. Gen. Distr. : 
probably occurring in adjacent parts of Turkey. Described from Armenia. 

Note. Apparently forming hybrids with S. c auc as ic a (E. Transc). 



Series 3. Euariae Zinserl.— Sepals persistent in mature fruits. Leaves 
unlobed, dentate or duplicato-denticulate, gray-tomentose below; teeth 
numerous (more than 20 on each side); lateral veins 7 — 12. 

Note. Species of this series, native to part of Bal. -As. Min., do not 
grow wild in the USSR. 

*S. aria Crantz, Stirp. austr. fasc. I (1762) 46. 

Tree; leaves rounded- elliptic to elliptic, ovate-elliptic, or obovate- elliptic, 
gray-tomentose below, with 8—12 pairs of lateral veins, usually cuneately taper- 
ing downward, obtuse or acuminate. Growing wild in Central Europe. 



Series 4. Graecae Zinserl. —Sepals persistent in mature fruits. Leaves 
dentate or coarsely duplicato-dentate, white-tomentose below (less often 
gray or gray-green-tomentose); teeth more than 20 on each side; lateral 
veins not more than 10 pairs. 



303 



398 



24. S. kusnetzovii Zinserl. sp. n. in Addenda VIII, p. 382. 

Tree; buds tomentose-pubescent, sometimes subglabrous; leaves broadly 
obovate-elliptic, fusiform-elliptic, or elliptic, cuneately tapering toward 
base, acuminate (less often subobtuse), 7 — 13 cm long, 4 — 9 cm broad (length- 
to-width ratio 1.4 — 1.8), with 8 — 10 pairs of lateral veins, the blade glabrous 
above (except for midrib), gray-tomentose or gray-green-tomentose below, 
(less often white-tomentose), the margin — except for lower % — conspicuously, 
less often obscurely duplicato-dentate, with 30 — 40 sharp teeth of the 
second order; pedicels white-pubescent; calyx white-tomentose, with 
acutely triangular teeth, more or less closing after anthesis; petals 
orbicular, clawed; fruits globose -ovoid (mature fruits unknown). Fl. 
June -July. (Plate XXVI, Figure 11). 

Rocks, pine forests, at 1,600 — 1,850 m.— Caucausus: Cisc, Dag., S. 
Transc. (Lake Sevan). Endemic. Described from the Caucasus. Type in 
Leningrad. 

Note. This species links the series Graecae to the series Euariae 
by its somewhat larger number of lateral veins and especially by the gray 
or gray-green color of underside of leaves. Specimens from Armenia are 
distinguished by the densely white-pubescent underside of their leaves. 

25. S. graeca (Spach) HedL, Monogr. d. Gatt. Sorbus (l90l) 75.— 

S. umbellata Fritsch in Sched. Fl. Exs. Austro-Hung. (1896).— S. um - 
bell at a var. graeca C. K. Schn., Handb. d. Laubholzk. I (1906) 689 p. p. — 
S. umbellata var. c r etica Grossg., Fl. Kavk. IV (1934) 2 88 p. p. — 
S. cretica Fritsch Sched. ad. Fl. exs. Austro-Hung. (1896). — S. ar i a var. 
graeca K. Koch, Dendr. I (1896) 102; Boiss., Fl. Or. II (l 872) 658 p. p. - 
Pirus aria y cretica Lindl. in Trans. Hort. Soc. London VII (l 828) 
235 p. p.— P. graeca Loddiges, Cat. (1816) 26 (nom.nud.).— Crataegus 
graeca Spach, Hist. nat. d. Veg. II (1834) 102. — Aria graeca Roem., 
Fam.nat. Syn. Ill (1847) 127. Ic: HedL, Monogr. d. Gatt. Sorbus (l 901 ) 
75, f. 20; C. K. Schn., Handb. d. Laubholzk. I, 690, f. 379 e— i. 

Small tree or large shrub; buds slightly tomentose or subglabrous; 
leaves coriaceous, orbicular or rounded-elliptic, rounded toward base 
(var. or biculata Zinserl.) or else obovate-elliptic or obovate, cuneately 
tapering toward base (var. cuneata Zinserl.), obtuse or acuminate, 
5 — 10 cm long, 4 — 10 cm broad (length-to-width ratio 1—1.3) with 7 — 9 (10) 
pairs of lateral veins, the blade glabrous above (except midrib) (pubescent 
when young), densely white-tomentose below throughout the lower surface 
including the veins, the margin — except for lower part — with inconspicuous 
lobes; teeth of the second order acute, 20 — 3 5 on each side; pedicels white- 
tomentose; petioles 0.3 —1.5 cm long, white-tomentose; calyx white- 
tomentose, with acute triangular teeth, more or less closing after anthesis; 
petals rounded-ovate; fruits globose, red, later turning blue. Fl. June — 
July, fr. August — September. (Plate XXVI, Figures 10 and 12). 

Rocks, taluses, forests in upper part of the forest zone and in the 
subalpine zone, at 1,200 — 2,500 m. — European part: Crim.; Caucasus: 
Cisc, W., E., and S. Transc, Tal. Gen. distr. : Bal. -As. Min., Centr. Eur. 
Described from Greece. Type in Paris. 

26. S. migarica Zinserl. sp. n. in Addenda VIII, p. 382. 

Shrub 30 — 50 cm high; buds slightly tomentose; bark dark gray-brown 
in young branches); leaves more or less coriaceous, orbicular, obtuse, 



304 



4 — 5 cm long, 3 — 5 cm broad (length-to-width ratio 1 — 1.2), with 8 — 10 pairs 
of veins, glabrous or slightly pubescent (except veins) above (pubescent 
when young), with conspicuously impressed veins, densely white-tomentose 
below between the veins, much less pubescent along the veins, hence the 
veins conspicuous by their darker coloration, the blade entire in lower 
%— l lz> otherwise dentate (in some leaves the dentation scarcely discernible), 
with 20 — 25 small, acute teeth on each side; petioles 0.2 — 0.4 cm long, 
tomentose; corymbs few-flowered; pedicels white-tomentose; calyx 
white-tomentose, with triangular teeth recurved after anthesis. Flowers and 
mature fruits unknown. Immature fruits in July. 

Limestone slopes at ca. 2,000 m. — Caucasus: W. Transc. (Mingrelia, on 
Mt. Migariya). Endemic. Described from the Caucasus. Type in 
Leningrad. 

27. S. baldaccii Deg. et Fritsch in herb. — S. m e r id i onal i s var. 
baldaccii Asch.et Gr. Synops. VI, 2 (1906) 100. - S. u mb e 1 1 a t a var. c. 
baldaccii C. K. Schn., Handb. d. Laubh. I (l 906) 689. - Ic. : C. K. Schn., 
Handb. d. Laubh. I, 690, f. 37 9 I-n. 
3 gg Small tree (?) or shrub (?); buds slightly tomentose; bark dark gray, 
brown on young branches; leaves more or less coriaceous, oblong- 
elliptic, to ovate-oblong-elliptic, obtuse or acuminate, cuneately tapering 
or rounded toward base, 5 — 7cm long, 3— 4cm broad (length-to-width ratio 
1.7 — 2.0), glabrous (except veins) above (pubescent when young), densely 
white-pubescent below, with 6 — 9 pairs of veins (7 — 9 in Caucasian-Asia 
Minor specimens), the blade — except lower entire part {%—%) — obscurely 
or conspicuously duplicato-dentate, the teeth of the second order acute, 
20 — 25 on each side; petioles 1— 1.5 cm long; corymbs few- to many- 
flowered; pedicels white-tomentose; calyx white-tomentose with triangular 
teeth, more or less closing after anthesis. Mature fruits unknown. Fl. 
July — August. 

Mountain slopes in the forest zone.— Caucasus: S. Transc. (Nakhichevan 
ASSR near Lake Batabad). Gen. distr. : Bal. -As. Min. (Albania, Cappadocia, 
parts of Turkish Armenia bordering on the USSR). Described from Albania. 
Type in Vienna. 

Note. Apparently forms hybrids with S. g r a e c a (Spach) Hedl. (near 
Lake Sevan in the Armenian SSR). It may be that Asia Minor forms differing 
from the Albanian specimens (with ca. 6 veins) should be separated as a 
species. 



Series 5. Xerophilae Zinserl. — Sepals persistent in mature fruits. 
Leaves dentate or duplicato-dentate, sometimes shallowly lobed (S. per sic a 
Hedl.), white-tomentose below, the teeth (simple or of the second order) 
not more than 20 on each side (35 only in S. p e r s i c a) Lateral veins 5 — 8 
pairs. 

2 8. S. turcica Zinserl. sp. n. in Addenda VIII, p. 383.- S. flabellifolia 
Hedl., Monogr. d. Gatt. Sorbus (1901) 71, non al. — S. umbellata var. 
flabellifolia C. K. Schn., I (1906) 689 p. p.; Grossg., Fl. Kavk. IV (1934) 
288.— S. aria var. graeca Boiss., Fl. Or. II (1872) 658, p. p. 



305 



400 



Shrub or small tree; buds slightly tomentose; leaves coriaceous, 
orbicular, obtuse or scarcely acuminate, rounded or slightly narrowed at 
base, 5 — 7 cm long, 4 — 6 cm broad (length-to-width ratio 1.1 — 1.3), with 
6 — 8 pairs of lateral veins, glabrous above (adult), densely white-tomentose 
below, the margin entire to A or higher, simply dentate or obscurely 
duplicato-dentate in upper part, the teeth large, abruptly narrowing from 
rounded base, acute, 10 — 1 5 on each side; petioles 0.5 — 1 .2 cm long, 
tomentose; flowers white-tomentose; calyx white-tomentose, with triangular 
teeth; fruits globose, red, turning blue- Fl. May, fr. (mature) September. 

Forests, rocks in lower parts of the forest zone.— European part: Crim.; 
Caucasus: S. Transc. Gen. distr. : Bal. -As. Min (As. Min. ). Described from 
Asia Minor. Type in Leningrad. 

29. S. taurica Zinserl. sp. n. in Addenda VIII, p. 383. — S. umbellata 
var.flabellif olia C. K. Schn., I (1906) 689 p. p. - Exs. : HFR No. 969 
p. p. (sub S. a r i a var. g r a e c a Boiss.). 

Shrub or tree; buds slightly tomentose; leaves coriaceous, obovate or 
obovate-elliptic, obtuse or slightly acuminate, conspicuously cuneately 
tapering toward base, 5 — 7 cm long, 3.5 — 5.5 cm broad (length-to-width ratio 
1.3 — 1 .5), with (6) 7 — 8 (9) pairs of lateral veins, glabrous above (adult), 
when young tomentose-pubescent, densely white-tomentose below, the 
margin entire to jz or more^rom base, in upper part dentate or 
obscurely duplicato-dentate, the teeth large, abruptly narrowing from 
rounded base, acute, 10 — 1 5 on each side; petioles 0.5 — 1.5 cm long, white- 
tomentose; pedicels white-tomentose; calyx white-tomentose, with 
triangular teeth; fruits globose, red, turning blue. Fl. May — June, fr. 
September. 

Rocks, oak and juniper forests.— European part: Crim.; Caucasus: 
W. Transc. (vicinity of Novorossiisk). Endemic. Described from the 
Crimea. Type in London. 

Note. S. t au r i c a, closely related to S. m e r i d i onal i s (Guss.)Nym.— 
distributed in the W. Med. (Sicily, Sardinia, possibly Crete) — is distinguished 
from the latter by its leaves: less sharply cuneate-tapering toward base, 
with less acute teeth. The species S. flabellifolia Schauer, to which 
S. taurica and S. turcica are often referred, is described after a 
cultivated specimen of imprecisely known origin, and is distinguished from 
those species by its lobate leaves. S. taurica and S. t u r c i c a, differing 
only in unimportant characters and having overlapping distribution areas, 
are not clearly delimited; in the Crimea there are very many specimens 
transitional between these two species, whereas Caucasian and Asia Minor 
specimens have clearly expressed S. taurica characters. Apparently this 
is a case of as yet uncompleted differentiation of one species into two — 
the Caucasian-Asia Minor S. turcica and the Crimean S. t a u r i c a. In the 
Crimea, S. taurica (possibly also S. t u r c i c a) often forms hybrids with 
S. g r a e c a. 

30. S. obtusidentata Zinserl. sp. n. in Addenda VIII, p. 384. 

Small tree (? ) or shrub (? ); buds glabrous; bark dark gray, brown on 
young branches; leaves rounded-elliptic or obovate, obtuse or slightly 
acuminate, pointed or attenuate at base, 6 — 9 cm long, 5.5 — 7 cm broad 
(length-to-width ratio 1.1 — 1.3), glabrous — except on the veins — above 



306 



(pubescent when young), densely white -tomentose below, with 6 or 7 pairs 
of veins, the blade entire or subentire in lower / 4 — 1$, dentate or else 
duplicato-dentate (and then teeth of the first order obtuse, crenate), the 
teeth of the second order subobtuse, small, 15 — 20 on each side; petioles 
0.3 — 0.5 cm long; pedicels white-tomentose; calyx white -tomentose; 
fruits unknown. July. 

Alpine zone at 2,250m.— Caucasus: W. Transc. (Abkhazia, Mingrelia). 
Endemic. Type in Leningrad. 

31 . S. persica Hedl., Monogr. d. Gatt. Sorbus (l 901 ) 70; C. K. Schneid., 
Handb. d. Laubh. I, 694; Grossg., PI. Kavk. IV (l 934) 288. — Ic : Hedl., 
Monogr. d. Gatt. Sorbus, 70, f. 18; C. K. Schn., 1. c, 690, f. 3790. 

Tree; buds slightly pubescent; leaves elliptic, rounded-elliptic, or 
oblong- elliptic, obtuse or subacute, slightly cuneately tapering toward base, 
5.5 — 7.5 (9.5) cmlong, 3.5 — 4.5 (6.5) cm broad (length-to-width ratio (1.1) 
1.4 — 1 .5 (1.7)), 4 — 6-lobed (lobes passing into crenae at the apex), depth of 
lower lobe and middle lobe %—% the half-width leaf, coriaceous, glabrous 
above, white-tomentose below, with 25 — 35 teeth on each margin, the teeth 
acute, obliquely triangular, larger in upper than in lower part; lateral 
veins 5 — 7 (8), forming with the midrib an angle of less than 45°; lobes 
oblong-rounded, less often oblong-triangular, obtuse, less often acuminate, 
with few teeth at outer margin, with 1 or 2 teeth on inner margin; petioles 
tomentose-pubescent, 1— 2 cm long; corymbs many-flowered; pedicels 
tomentose at anthesis, later glabrous; calyx with acute triangular teeth, 
tomentose; petals white, rounded-obovate, strongly narrowing toward base; 
fruits globose, less often ovoid, bluish red. Fl. June, fr. September. 

Mountains, shrubs, juniper and deciduous forests, 1,300 — 2,800 m (rarely 
down to 850 m).— Caucasus: S. Transc. (very rarely); Centr. Asia: Mtn. 
Turkm., Pam. -Al., T. Sh. Gen. distr. : Iran. Described from N. Iran. 

SPECIES INTERMEDIATE BETWEEN THE SECTION ARIA 

(SUBGENUS HAHNIA) AND THE SUBGENUS 

EU-SORBUS 

These species are characterized by leaves pinnatifid in lower part and 
by styles usually free or nearly free. Apparently all or at least most of 
then are the result of crossing between species of the section Aucuparia 
and Aria, producing a stable progeny. 



32. S. turkestanica (Franchet) Hedl., Monogr. d. Gatt. Sorbus (1901) 69; 
C. K. Schn., 111. Handb. I, 692. — Pi rus turkestanica Franchet, PI. turk. 
in Ann. d. Sc. Nat. Ser. VI, bot. (1883) 288.— Ic: C. K. Schn., I.e., f. 379, u. 

Tree; buds slightly pubescent; leaves oblong- elliptic, acute or subobtuse, 
cuneately tapering toward base, (5.5) 6 — 10 (12) long, (3) 3.5 — 6.5 (8)cm 
broad (length-to-width ratio 1 .5 — 1 .7), pinnatisect, with 1 or 2 pairs of 
segments in lower part, in greater part of the tree lobate, pinnatifid, the 
depth of lower and middle lobes % — % the half-width of blade, coriaceous, 
glabrous above, gray-tomentose below, with 30 — 50 teeth on each margin, 
the teeth small, obliquely triangular-lanceolate, acute, sometimes subaristate; 
lateral veins 7 — 9, forming an angle of 45° with the midrib; lobes oblong 
or oblong-triangular, acute, less often subobtuse, with few teeth on outer 



307 



405 



margin and with 1 or 2 minute teeth in inner margin; petioles tomentose- 
pubescent, 0.7 — 2 cm long; corymbs many-flowered; pedicels glabrous at 
anthesis (less often slightly pubescent); receptacle glabrous; calyx 
tomentose, with narrowly lanceolate teeth; petals white, rounded-obovate, 
tapering toward base; fruits ovoid or ovoid-pyriform, turning blue. 
Fl. May — June, fr. September. 

Mountains in forests and among shrubs, at 1,150 — 2,600 m. — Centr. Asia: 
Pam. -AL, T. Sh., Mtn. Turkm. (? ). Endemic. Described from Central 
Asia. Type in Paris. 

Note. This species may be the result of a cross between S. persica 
Hedl. and S. tianschanica Rupr. 

33. S. dualis Zinserl. in Addenda VIII, p. 384. 

Tree; buds slightly tomentose (sometimes only scale margins); leaves 
ovate or oblong-ovate, obtuse or acuminate, 6 — 10cm long, 4.5 — 7.5 cm 
broad (length-to-width ratio 1.2 — 1.6), pinnatisect in lower part, with 
1—3 pairs of segments, gradually passing into pinnatifid higher up, with 
8 — 11 pairs of lateral veins, gray-green and glabrous (adult) above except 
midrib, gray-tomentose below, the segments and leaf lobes dentate- 
margined, the teeth acute, passing abruptly from rounded base into a usually 
bent point, numerous (40 — 50 or more on each side of blade), the segments 
and lobes with teeth on both margins; petioles 1—2 cm long, tomentose; 
pedicels tomentose, glabrous in mature fruits; calyx with acute -triangular, 
more or less tomentose teeth persistent in mature fruits; fruits globose, 
red, later turning blue. Fl. unknown; fr. (mature) September. 

Forests (especially of Quercus macranthera) to 1,500m.— 
Caucasus: E. and S. Transc. Described from the Caucasus. Type in 
Leningrad. 

Note 1. This species is apparently the result of a cross between 
S. armenica Hedl. (which it resembles in character of teeth) and 
S. aucuparia L. In any case, it is undoubtedly the result of crossing 
between species of the sections Aucuparia and A r i a . 

Note 2. In regions where species of the sections Aria and Aucu- 
paria grow together there occur hybrids characterized by their leaves, 
which are pinnatisect at the base, lobate higher up. Such hybrids have been 
found in W., E., and S. Transc. Some of these hybrids are cultivated. 

♦S.fennica (Kalm. ) Fr., Summaveg. Scand. (1846) 42.— Crataegus 
fennica Kalm., Fl. fenn. I (1765) 6. —Distinguished from the preceding 
by its oblong or oblong-ovate leaves with somewhat less acute teeth. 

Sometimes cultivated in parks. Grows wild in Scandinavia and the 
Baltic States. According to Hedlund, it is a hybrid of S. aucuparia and 
S. salicifolia Hedl. 



Section 2. TORMINARIA DC, Prodr. II (1825) 636.- Flesh of fruit with 
numerous grit cells; carpels almost completely connate. 

34. S. torminalis (L.) Crantz, Stirp. austr. II (1767) 45.— Crataegus 
torminalis L., Sp. pi. (1753) 46. — Py r u s torminalis L., Sp. pi. 
(1753) 46.— Pirus torminalis Ehrh., Beitr. Naturk. IV (l 789) 92. — 
Hahnia torminalis Medik., Gesch. Bot. (1793) 81. 



308 



403 




PLATE XXVI. 1— Sorbus schem a che ns i s Zinserl., leaf; 2— S.b aid ac ci i Deg.et 
Fritsch, leaf; 3-S.migarica Zinserl., leaf; 4-S.torminal is Crantz, flowering branch; 
5-S. turcica Zinserl., leaf; 6-S.taurica Zinzerl., leaf; 7 -S.ob tu si d en t a t a Zinserl., 
leaf; 8-S.buschiana Zinserl., leaf on sterile shoot; 9-S.buschi an a Zinserl., leaf;on 
fruiting shoot; 10-S.graeca var.o rb icul at a Zinserl., leaf; 11 -S.kusne t so vii Zinserl. 
leaf; 12— S.graeca var.cuneata Zinserl., leaf. 



309 



406 



Tree, usually to 12 m, sometimes to 25 m high, to 60 cm in diameter 
(usually less); buds glabrous; leaves ovate, rounded or slightly cordate or 
cuneate at base, acute, 5 — 10 cm broad, 16 — 18 cm long, with 3 — 5 acute 
(less often subobtuse) lobes, sometimes the lobes — especially the lowest 
pair — very acute and deep, since leaves nearly pinnatifid at base (var. 
pinnatifida Boiss.), acutely denticulate, glabrous above (pubescent only 
when young), yellow-green and glabrous below (pubescent when young) or 
slightly tomentose (var. mollis Beck), with 3 — 7 pairs of lateral veins; 
petioles 2 — 5 cm long; pedicels initially villous -tomentose, later glabrescent; 
calyx more or less tomentose, with triangular teeth; petals white, spreading; 
styles 2; fruits ovoid or globose, brown to yellow with white dots. May. 

Forests, mainly oak but also chestnut, hornbeam and Pinus nigra, 
from plains to 1,700 m. —European part. : M. Dnp., Bl., Crim. ; Caucasus: 
Cisc, W., E., and S. Transc, Tal. Gen. distr.: Atl. and N. Eur., Scand., Bal. - 
As. Min. Described from Central Europe. Type in London. 

Economic importance. Fine-grained, heavy wood (specific gravity 0.8) 
responding well to polishing, valuable for turning and carving. Fruit edible. 
Ornamental plant. 

Note. Forms hybrids with species of the section Aria. Similar plants 
with less deeply-lobed leaves and more obtuse lobes gray-tomentose below 
have been found in fruit in the Crimea. 



Genus 729. MICROMELES * DECNE.** 

Decne. in Nouv. Arch. Mus. Paris X (1874) 168. 

Flowers small, mostly in threes in umbelliform corymbs, the latter in 
groups at ends of shoots. Ovary of 2 or 3 (4) carpels (styles 2 or 3, 
rarely 4), connate and adnate to the hypanthium to lz — h- Pome very 
compact, without nutlet but with 2 — 4 seeds; scales not persistent in fruit. 
Trees and shrubs with leaves caducous or persistent in winter, simple, 
dentate. Approximately 10 species in the Far East and SE Asia. 

1. M. alnifolia (Sieb. et Zucc.) Koehne, Die Gattung. Pomac. (1890) 20; 
Kom., Fl. Manch II (1904) 479; Kom. and Alis., Opred. rast. Dal'nevost. 
kraya II (1932) 638.— Crataegus alnifolia Sieb. et Zucc. in Abh. Acad. 
Wissensch. Munch. IV, 2 (1846) 130.— Sorbus alnifolia C. Koch in Ann. 
Mus. bot. Lugd. Bat. I (1863) 249.— Aria alnifolia Decne in Nouv. Arch. 
Mus. Paris X (1874) 166.— Pyrus miyabei Sarg. in Gard. a. For. (l 893) 
214.— Sorbus miyabei Mayr, Fremdl. Waldb. (l 906) 491 . — Ic. : Kom. 
and Alis., I.e., Plate 191, Fig. 3.— HFR No. 1323. 

Slender tree to 18m high, forming a rather sparse crown only high above 
the ground, with straight branches and glabrous or initially hairy shoots; 
leaves mostly ovate and short-acuminate or oval and more or less long- 
acuminate, rounded or rounded-cuneate at base (var. t y p i c a C. K. Schn. ) 
less often orbicular with cordate base (var. tiliifolia (Decne.) C. K. Schn. ), 
unequally duplicato-dentate, dark green and lustrous above, slightly hairy 
only along the veins above, yellowish-hairy along the veins below (when 
young, sometimes with continuous pubescence) or glabrous; flowers ca. 1 cm in 



From the Greek micros, small and melon, pome. 
Treatment by A.I.Poyarkova. 

310 



diameter, 2 or 3 in each long-stalked, umbelliform corymb, the latter in 
groups at ends of lateral branches; sepals white-tomentose inside; fruits 
very rigid, dry, oblong, 8 — 12 mm long, 6 mm broad, with glaucous bloom. 
Fl. June, fr. from August. v Plate XXVII, Figure 2). 

Shady mountain forests of Siberian stone pine and deciduous trees, 
slopes, on stony and humous soils.— Far East: Uss. Gen. distr. : Jap. -Ch. 
planch., Korea, S. China and Japan). Described from Japan. 

Economic importance: The wood is characterized by its even grain and 
is used for making rulers. Fruits edible. 



Genus *ERIOBOTRYA* LINDL. ** 
Lindl. Trans. Linn. Soc. XIII fl821) 102. 

Flowers in pyramidal panicles; ovary of 5 carpels (styles 5), completely 
connate, overgrown by the hypanthium except at the apex; carpels 2-ovuled. 
Pome succulent, with very thin coriaceous endocarp and large globose - 
angular seeds. Trees or shrubs with entire evergreen leaves.— 
Approximately 10 species in SE Asia. 

1. E. japonica Lindl. in Trans. Linn. Soc. XIII (1821) 102.— Mes pilus 

japonica Thunb., Fl. jap. (1 784) 206. — Pho t i n i a japonica Fr. et Sav.. 
PI. jap. (1875) 142.— Ic: Bot. Reg. tab. 365; Hegi, Illustr. Fl. v. Mitteleur. 
IV, 2 (192 5) f. 998. 

Small tree or shrub, with dense rufous-gray tomentose pubescence 
covering shoots, underside of leaves, and inflorescence; leaves rigid - 
coriaceous, evergreen, glabrous above, dark green, large, to 2 5 cm long 
and 7 — 8 cm broad, lanceolate or oblong-lanceolate, acute, with decurrent 
cuneate base, minute-petioled, mostly with coarse, sparse teeth, sometimes 
only in upper part, some leaves entire; inflorescence a broadly pyramidal 
panicle to 17 cm long; flowers inconspicuous owing to dense pubescence 
COA ering all parts of inflorescence, aromatic; sepals short, rounded- obovate; 
petals white, pubescent inside; stamens 20; fruits large, ca. 3 cm in 
diameter, with persistent sepals at the apex, glabrous at maturity, yellow, 
3 — 5-locular, with few, mostly 2 or 3, less often 1 or 4 — 7 large seeds, very 
succulent, bittersweet. Fl. March, fr. April — June. i^Plate XXVII, Figure 3), 

Cultivated in the Crimea and the Caucasus, growing wild in W. China 
and Japan (? ). Cultivated since ancient times in India, Japan, and China. 
Introduced into cultivation in Europe in 17 88. Described from Japan. 
Type in London. 

Economic importance. Cultivated as an ornamental for its splendid 
foliage and tasty fruit. The ripe fruits contain 6 % invert sugars, 4. 94 % 
saccharose. 0.6 % malic acid ^no other acids), 3% pentazans. The seeds 
contain a small quantity of prussu- acid, amygdalin, laurocerasin, and saponin. 



From the Greek eri 6s, hairy and botrys, raceme, 
Treatment by A.I.Poyarkova, 



311 



409 



Genus 730. AMELANCHIER * MEDIK.** 

Medik., Phil. bot. I (1789) 155. 

Flowers small, white, in racemes, with (15) 20 stamens; ovary usually 
of 5 (2 — 5) carpels connate only at base, sunken to the middle in the 
hypanthium, each ovary separated into two 1-ovuled locules by an 
incomplete (false) partion. Pome succulent, without nutlets, the endocarp 
membranous. Shrubs or trees with simple, entire leaves. 

Economic importance. All Amelanchier species have edible fruit 
and wood suitable for making small articles; most species are valued as 
ornamentals. 

1. Leaves entire *2. A. integrifolia Boiss. et Hob. 

+ Leaves dentate 2. 

2. Styles free, scarcely attaining base of stamens 

1. A. rotundifolia (Lam. ) Dum. -Cours. 

+ Styles fused, at least below, attaining anthers of the inner series 

of stamens 3. 

3. Ovary apex glabrous; fruits dark purple with glaucous bloom; 
sepals reflexed *A. canadensis (L. ) Medik. 

+ Ovary apex tomentose-hairy 4. 

4. Sepals recurved *A. turkestanica Litw. 

+ Sepals erect; fruits black-glaucous A. spicata (Lam. ) C.Koch. 

1. A. rotundifolia (Lam.) Dum. -Cours., Bot. Cult. ed. 2, V (1811 ) 459; 
C K. Schn.,Ill.Handb.d. Laubh. I (1906) 732; Grossg., Fl. Kavk. IV (1934) 
289.— Mespilus amelanchier L., Sp. pi. (l 753) 478. — Py r u s ame- 
lanchier Willd., Sp. pi. ed. 2 (1798) 1014; M. B., Fl. taur. -cauc. I (1808) 
389.— Sorbus amelanchier Crantz, Stirp. austr. I, II (1762) 53. — 
Crataegus rotundifolia Lam., Encycl. meth. I (l 783) 83.- Ame- 
lanchier ovalis Medik., Gesch. bot. (1793) 79. — A. vu lg a r i s Moench, 
Meth. (1794) 682; Ldb., Fl. Ross. II, 1, 93; Shmal'g. I, 353; Medv., Der. i kust. 
Kavk. (1919) 117.— Aronia rotundifolia Pers., Synops. II (l 807) 39. - 
Crataegus amelanchier Desf., Hist. arb. I (l 809) 149. — A m e 1 an - 
chier rupestris Bluff et Fingerh., Comp. fl. Germ. I (l 825) 609. — Ic. : 
Vol'f and Palib., Opred. der. i kust. (1904) figures on pp. 433 and 434; Hegi, 
Illustr. Fl. v. Mitteleur. IV, 2 (1925) tab. 146, f. 3 -3a, textfig. 1073-1076. 

Shrub 0.5 — 2 m high; shoots whitish-tomentose when young, glabrous 
later; leaves firm, elliptic, ovate, or obovate, rounded at the apex, often 
emarginate, less often very short-acuminate, mostly rounded or slightly 
cordate at base, continuously whitish-flocculose below when young but 
glabrous when g]abrous above initially, with simple, acute, antrorse teeth 
on the margin; adult stipules long, linear, caducous: flowers in dense, cory- 
mbiform, 5 — 8-flowered racemes; pedicels of upper flowers 2 — 5 mm long, 
those of lower flowers not more than 10 — 12 mm long, initially tomentose 
like the hypanthium, later glabrous; sepals triangular-lanceolate, acuminate, 
glabrous or initially tomentose, erect in fruit; petals oblong-lanceolate to 
linear, slightly hairy exteriorly, 13 — 1 6 mm long, 2 — 5 mm broad; stamens 
20; pistils 2 — 5; ovary apex tomentose; styles very short, not protruding 
from the hypanthium, free; fruits the size of a pea, succulent, initially red, 

* Apparently from the Provencal amelanche .referring to the honey taste of the fruit. 
** Treatment by A.I.Poyarkova. 

312 



at maturity black with glaucous bloom. Fl. May, fr. from end of 
August. (Plate XXVII, Figure 4). 

Middle mountain zone, to 1,900 m, rocky sites among shrubs, forest 
edges and open woods. — European part: Crim.; Caucasus: Cisc, Dag., 
W., S., and E. Transc; Gen. distr.; Centr. and S. Eur., As. Min., S. Afr. 
Described from Switzerland. Type in London. 

Economic importance. Beautiful shrub cultivated in the South as an 
ornamental plant, used for hedgerows; sweet fruit edible. Wood fine- 
grained, reddish white, hard, very heavy, used for making small articles. 
Reproduces by seeds or suckers. 

*2. A. integrifoliaBoiss, etHoh. in Boiss., Diagn, ser. I, III (1843) 
8.— A. vulgaris var. integrifolia Boiss., Fl. Or. II (1872) 667.— Ic: 
CK. Schn.,Ill.Handb. Laubh. I (1906) f. 407 e-f, 408 e-f. 

Shrub; leaves entire, only single leaves dentate near the apex, more 
coriaceous than in the preceding species, oval, rounded on both sides, 
mucronulate, less often short- acuminate or emarginate, glabrous above, 
below covered like the shoots with dense tomentum persisting in its greater 
part until fruiting (always ? ); flowers in sparse 4 — 6 -flowered racemes; 
pedicels of upper flowers 5 — 10 mm long, those of lower flowers to 20 mm, 
their tomentose pubescence dense and long-persistent as on the hypanthium 
and sepals; sepals triangular, erect in fruit; petals oboval, 8 — 10 mm 
long, 4 mm broad, hairy outside; fruits as in the preceding species. 

Rocky sites, not higher than the middle forest zone. Not yet found in the Soviet 
Union — nearest site in the Kagyzman district. Gen. distr.: Bal. -As. Min. 
(Galacia), Arm. -Kurd. Described from Mt. Gara in Kurdistan. Type 
in Geneva; cotype in Leningrad. 

Economic importance. Ornamental. Fruits edible. 

3. A. turkestanica Litw. in Trav. Mus. bot. Acad. Sc. St. Petersb. VIII 
(l 91 1 ) 80.— A. asiatica var. turkestanica Litw., 1. c, p. 7 9; Fedchenko 
Rast. Turk. (1915). 

Shrub; leaves ovate, obtuse, cordate at base, with acute teeth from the 
middle to the apex, 2—4 cm long, 1.5 — 2.5 cm broad, glabrous or with 
appressed hairs in some places; petioles to 2 cm long, scattered-hairy; 
sepals triangular, recurved, glabrous outside, hairy inside; petals to 9 mm 
long, lanceolate-cuneate, subobtuse, partly scattered-hairy outside; 
styles 5, connate to /j, sparsely hairy at base, not exceeding inner series 
of stamens. 

Described after a specimen allegedly collected on stony hills in the 
Bayanaul Mountains in the Semipalatinsk Region. Type in Leningrad. 

Note. The finding of an Amelanchier specimen, of North American 
parentage according to all its characters, in a region so remote from its 
distribution area obviously requires verification. Has there not been a 
confusion of labels? 

*A. canadensis (L. ) Medik., Gesch. Bot. (1793) 79. — M e s p ilu s cana- 
densis L., Sp.pl. (1753) 478.-A. botryapium DC., Prodr. II (182 5) 632.- 
Ic: Britt. a. Brown., 111. Fl. N. Amer. II (1913) figure on p. 292; C. K. Schneid., 
Ill.Handb. Laubh. I (1906) f. 409a- c, 410a. 



313 



411 




PLATE XXVII. 1— Oydonia oblonga Mill.: a) section of fruit; 2 — Micromeles alnifolia 
Koehne; 3 — Eriobotrya japonica Lindl. ; a) fruit; 4— A mel anchi er rotund if oli a 
(Lam.) Dum. -Cours. ; — M espi 1 us germanica L. : a) fruit. 



314 



Shrub or tree, usually 7 m, sometimes to 18 m high; leaves oval, acute 
or acuminate, thin, covered below in frondescence with soon disappearing 
rufous tomentum, rounded or cordate at base, serrulate; flowers in erect or 
slightly nodding rather sparse racemes; pedicels long, from 6 mm (the 
upper) to 2.5 mm (the lower), axis and pedicels initially hairy, later glabrous; 
hypanthium and lanceolate-triangular sepals usually devoid of pubescence 
from the very beginning; petals oblong, linear or spatulate; ovary apex 
glabrous; styles connate to the middle; fruits dark purple with glaucous 
bloom, with reflexed sepals at the apex. 

Often cultivated in European part of Soviet Union; originally from 
N. America, where it grows in dry deciduous forests. Described from 
N. America. Type in London. 

Economic importance. Ornamental plants; fruit pleasant -tasting. Wood 
very hard, light brown, suitable for making small articles. 

*A. spicata(Lam. )C Koch, Dendr. I (1869) 182. — C r a t a egu s spicata 
Lam., Encycl. meth. I (1783) 84.— A. oval is Borkh., Forsbl. II (1803) 259 
(non Medik.).— A. c anad ens is var. spicata Sarg., Silva IV (l 892) 129. — 
Ic: C.K. Schn. I (1906) 411 m-n, 412 g — h.— Exs. : HFR No. 1019; PI. 
Pol. exs. No. 948 (sub A.botryapium Ser.). 

Tree or shrub to 5 m high; leave broadly elliptic, ovate, or oval, rounded 
at the apex or short-acuminate, initially tomentose below, later glabrous, 
acutely dentate; racemes dense, short, at blossoming sometimes densely- 
tomentose, later glabrous; sepals acute, oblong-triangular; petals 
8 — 12 mm long, oblong-oboval; ovary apex hairy; styles connate to 
about the middle, more than half protruding from the hypanthium; fruits 
black-glaucous, crowned by erect sepals. 

Very common in cultivation in central and NW regions of the European 
part of the Soviet Union; native to Canada and northern states of the U. S. A. 
Described from Canada. Type in Paris. 

Note. A.florida Lindl. (Bot. Reg. 1833, tab. 1 589) is also to be found 
in gardens of the Soviet Union: leaves with coarse teeth usually only from 
the apex to the middle, and sepals reflexed in glaucous -black fruits. 

Economic importance. Ornamental plants yielding edible fruit. 



Genus 731. PYRACANTHA * ROEM.** 

Roem.,Fam.nat.Syn. Ill (1847) 219. 

Flowers small, in compound, many-flowered corymbs, stamens 20; 
ovary of 5 carpels (styles 5), connate only at base; pome small, mealy, 
with 5 nutlets (endocarp stony), protruding to V3 — % from the flesh and 
covered by persistent sepals. Shrubs with entire leaves. Apart from the 
Soviet species 6 others are known, of which one in Formosa, the others 
in China. 

414 1. P. coccinea Roem., Syn. mon. Ill (1847) 219; Grossg., Fl. Kavk. IV 
(1934) 284.- Mespilus p y r ac an th a L., Sp. pi. (1753) 478; M. B., Fl. 
taur. -cauc. I, 388; III, 332. — C r at a eg u s pyracantha Medik., Gesch. 



* From the Greek pyr, fire and a ca nthos, thorn. 
** Treatment by A.I.Poyarkova. 



315 



415 



d. Bot. (1793) 84; Ldb., Fl. Ross. II, 1, 87. — C o t on e a s t e r pyracantha 
Spach, Hist. Nat. veg. II (1834) 73; Shmal'g., Fl. I, 348.- Ic: Pall., Fl. 
Ross. I, 1 (1784) tab. 13, f. 2; Vol'f i Palib., Opred. der. i kust. (1904) 
figures on pp. 441 and 442; Hegi, 111. Fl. Mitteleur. IV, 2 (1925) f. 1023, f . 
1025.— Exs.: HFRNo.1763; Callier, Iter taur. tert. No. 785. 

Shrub to 1.5 (2)m high, with broad, spreading crown, shoots grayish-hairy 
when young; branches dark reddish brown, covered with numerous spines, 
of which some short, 5 — 25 mm, leafless, others long, leafy, often with 
flowering shoots; leaves persistent, coriaceous, glabrous or else hairy 
only when young, dark green and lustrous above, lighter below, elliptic - 
lanceolate to narrowly lanceolate and tapering to short petiole or else ovate 
to ovate-lanceolate, with rounded base, with admixture of narrower leaves; 
all leaves acuminate, acute or obtuse to rounded at the apex, crenate, toward 
base often only undulate or even entire; inflorescence dense, more or less 
hairy; flowers small, white or pinkish yellow, with infundibular hypanthium; 
sepals broadly triangular; anthers red; fruits the size of peas, coral-red. 
Fl. June, fr. from September. 

Forest edges, slopes, among shrubs, to 1,200 m. — European part: Crim.; 
Caucasus: W., E., and S. Transc. Gen. distr.: Italy (possibly also SE France), 
Dalmatia, As. Min., N. Iran. Described from France. Type in London. 

Economic importance. Very ornamental shurb, owing to abundant 
blooming and beautiful long-persistent fruits. The latter are eaten by birds 
in winter. Used for hedgerows. Nectariferous: yielding nectar and pollen. 



Genus 732. MESPILUS L.* 

L.,Sp.Pl.(1753) 478. 

Flowers large, solitary; sepals exceeding petals; stamens 30 — 40; 
ovary of (4) 5 connate carpels adnate to the hypanthium; each carpel with 
2 ovules, only one of which develops; fruit a pome with (4) 5 nutlets 
(endocarp stony) entirely sunken in flesh. Spiny shrub or tree with simple, 
entire or dentate leaves. One species. 

1. M. germanica L. Sp.pl. (17 53)478; M.B., Fl. taur. -cauc, I, 388; Ldb., 
Fl. Ross. II, 1, 93; Shmal'g., Fl. I, 349. — M. c o mmun i s Giildenst., Iter I 
(17 87) 419, 42 8.- M. vulgaris Giildenst., 1. c, 312. — Ic. : Pall., Fl. Ross. 
I, 1 (1784)tab. 13, f. 1; Vol'f and Palib., Opred. der. i kust. (1904) fig. onp. 458; 
Hegi, Illustr. fl. v. Mitteleur. IV, 2 (192 5) tab. 146, f . 4, 4a, textfig. 1063, 1064.— 
Exs.:HFR No. 713 and 713a; Herb. Fl. Cauc. No. 326. 

Spiny shrub or small tree 1.5 — 6 m high, to 20 cm in diameter, with red- 
brown shoots pubescent when young, and with gray branches; leaves 
elliptic or oblong-lanceolate (3) — 5 — 12 cm long, acute or obtuse, entire 
or dentate with teeth terminating in a red gland, pubescent on both sides 
when young, later the upper surface dark green, scattered-hairy, more 
densely so along the veins, or else subglabrous, the underside light, whitish- 
pubescent, the longer and denser pubescence along main veins; petioles 
villous -pubescent; bracts large, caducous; flowers subsessile, white, 



Treatment by A.I.Poyarkova. 



316 



3 — 5 cm in diameter; sepals lanceolate -subulate, longer than petals; anthers 
purple, 1.5 — 2.5 cm in diameter, flattened-globose to pyriform, brownish, 
glabrescent. Fl. May, fr. October. (Plate XXVII, Figure 5). 

Up to 1,200 m, mainly at forest edges on slopes among shrub thickets, 
roadsides, clearings, also in forests on sufficiently moistened soils. 
Calcium-resistant. — European part: Crim. ; Caucasus: all districts; in 
Centr. Asia one site is known in the Kopet Dagh Range in the Gyuen Gorge 
on the Chandyr River. Gen. distr.: S. and SE Asia Minor, N. Iran. 
Described from S. Europe. Type in London. 

Note. Reports for the Caucasus on Mespilus smithii Ser. ex 
DC, Prodr. 11(1825) 633; Ldb., Fl. Ross. II, 1, 94; Medv., Der. i kust. 
(1919) 119 = Cratae-mespilus grandiflora (Sm.) Camus in Journ. 
d. bot. (1899) 326 = M. g e rmanic a L. X Crataegus monogyna 
Jacq., are unreliable, being based on leaves of sterile shoots which, unlike 
the typical Mespilus leaves, are slightly lobately incised. 

Economic importance. The fruits are edible, widely used by local 
population; the small wild forms as well as the large cultivated varieties 
are marketed. Owing to their tartness, the fresh fruits are used only after 
being affected by frost or when, after a period of storage, they become 
softer, more succulent and sweeter on fermentation. They make good 
marmalade and can be used for the making of drinks. 

The leaves, bark, and young fruits contain many tannins and are suitable 
for the tanning of leather. The fruit flesh contains 10.57 % invert sugars, 
5.84 % other nitrogen-free extracts, 7.51 % fibrous substances, 1.38 % malic 
acid; the seeds contain 2.5% fatty oils. Wood dense, very hard and heavy, 
yellowish white with reddish-brown heartwood, suitable for various turning 
jobs; produces coal of excellent quality, used as stock for grafting 
Japanese Mespilus. Ornamental. In the Soviet Union cultivated in the 
Crimea and in the Caucasus. There is no cultivated Mespilus in 
Central Asia. Among the cultivated varieties, the stoneless F. apyrena 
Duh. (f. abortiva Dum. -Cours) should be noted. 



Genus 733. CRATAEGUS * L.** 

L.,Sp.pl.(1753) 475. 

Flowers small, white, in mostly compound corymbiform, rarely simple 
umbellate inflorescences; ovary of 1 — 5 carpels free on ventral side or 
connate at base, adnate by the back to the hypanthium; ovules 2, the upper 
not developing; pome containing 1—5 nutlets (endocarp stony), with dry, 
persistent sepals at the apex, yellow, red, or black, rather fleshy. Small 
trees or shrubs with alternate stipulate pinnatilobate or partite dentate 
leaves. 

Economic importance. Ornamental trees and shrubs; many species are 
used for spiny hedgerows. The flowers are nectariferous and contain 
the yellow dye quercetin; their unpleasant odor is due to their trimethy- 
lamine content. The fruits of some species are large, fleshy, and tasty 



From the Greek k ra t o s , strength, sturdiness and a g e i n, to lead, to act, referring to the sturdiness and 
hardness of the wood or to the presence of hard thorns providing a means of defense. 
Treatment by A.I.Poyarkova. 



317 



417 



and are eaten, mostly raw, by the local population, mainly after frost; 
sometimes the fruit flesh is cut into small pieces and added to flour for 
the baking of sweet bread; sometimes the fruits are used for fruit jelly or 
stewed fruit or are even candied. The fruits are also eaten by birds, 
especially in Europe in winter, and are used as fodder for swine. 

The young leaves of certain species are used as a substitute for tea and 
are pleasant and refreshing to the taste. A decoction of leaves, bark, and 
roots is used for dying cloth light brown or yellow. 

The wood is fine-grained, dense, very hard and strong, heavy, reddish or 
yellowish, but difficult to split and treat; used for various lathework, 
tool handles, sticks, etc. 

Certain species are used in gardening as wildings for the grafting of 
apples, quince, and Japanese Mespilus. They reproduce by seeds which 
germinate with difficulty and are needed in stratification; very undemanding 
as far as soil is concerned. Since C r at a eg u s is readily attacked by various 
fruit tree pests, its presence near gardens is undesirable. 

1. Styles 3 to 5; fruits with 3 — 5 nutlets 2. 

+ Styles mostly 2, rarely 1 or 3, or else mostly 1; fruits with 2 

(1-3) nutlets 16. 

2. Inflorescence axes, pedicels, hypanthium, and outer side of sepals 
tomentose-pubescent; shoots and leaves — at least below — densely 
pubescent 3. 

+ Inflorescences glabrous, less often axes and pedicels slightly hairy, 
sometimes lower part of hypanthium rather densely pubescent; 
leaves glabrous or rather sparsely puberulent (not tomentosel). ... 7. 

3. Petioles % ( 2 /s _ lz) as long as the blade; inflorescence loose, with 
developed axes and pedicels 5. 

+ Petioles /s — % as long as the blade; inflorescence very compact, 

with short axes and pedicels (Series Orient ales) 4. 

4. Fruits with 5, rarely 4 nutlets, red-orange; sepals reflexed, 
lanceolate-triangular, long-acuminate; leaf lobes narrow: 3—4:1 
10. C. orientalis Pall. 

+ Fruits with 3 or 4 nutlets, (red?); sepals erect-spreading, broadly 
triangular, mucronulate; leaf lobes broader: 1.5 — 2.5 (3): 1 . . . . 
11. C. szovitsii A. Pojark. 

5. Leaves shallowly lobed; fruits red with with strongly pitted nutlets 
5. C. maxim oviczii C. K. Schn. 

+ Leaves partite; fruits black or cherry-red, with nearly smooth 

nutlets 6. 

6. Fruits black; inflorescence many-flowered, loose, large 

8. C. pentagyna W. et K. 

+ Fruits cherry-red; inflorescence few-flowered, small, rather 

compact 36. C. schraderiana Ldb. 

7. Leaves partite 8 

+ Leaves shallowly lobed (sometimes only with deep lower notches) 

12 

8. Fruits red or yellow to orange-brown 10 

+ Fruits black 9, 



318 



9. Corolla 10 — 12 mm in diameter; fruits without glaucous bloom; 
leaf notches usually narrow, i.e., lobes crowded; spines to 2 cm 
long 9- C. pseudomelanocarpa M. Pop. 

+ Corolla 13 — 17 mm in diameter; fruits with glaucous bloom, leaf 

notches mostly broadly triangular; spines not more than 12 mm long 

8. C. pentagyna W. et K. 

10. Fruits yellow to orange-brown 4. C altaica var. incisa Rgl. 

+ Fruits red 11- 

,11. Fruits small, 5 — 6 (8) mm long, mealy, with strongly pitted nutlets 

6. C remotibobata H. Raik. 

Fruits large, to 1.5 cm, with dense flesh and smooth nutlets 
l.C. pinnatifida Bge. 

12. Fruits black 13. 

+ Fruits yellow, orange-brown or red 14. 

13. Flesh of fruit green; nutlets strongly pitted; leaves usually with 

6 — 8 pairs of lobes 7. C. chlorosarca Maxim. 

+ Flesh of fruit yellow; nutlets smooth; leaves usually with 3 or 4 

pairs of lobes 33. C dsungarica Zbl. 

14. Fruits yellow or orange-brown, with 5 (4) nutlets; anthers white; 
leaves sometimes deeply incised 4. C. altaica Lge.* 

+ Fruits blood-red or orange -red, with 3 or 4 nutlets; anthers purple 

15. 

15. Fruits blood-red, mostly with 3 nutlets; leaves usually broadly 
obovate, puberulent on both sides 2. C. sanguinea Pall. 

+ Fruits lighter, to orange-red, with 3 or 4 nutlets; leaves mostly 

oblong -rhomboid or oblong -obovate, usually glabrous 

3. C. dahurica Koehne. 

16. Leaves, at least those on sterile shoots very deeply parted or almost 
dissected, with incised lobes; styles (nutlets) 1 or 2; shoots with 
numerous spines. (Series Stevenianae) 17. 

+ Leaves less incised 18. 

17. Mature fruits black 27. C. beckeriana A. Pojark. 

+ Mature fruits red 26. C. stevenii A. Pojark. 

18. Nutlet (style) mostly 1 19. 

+ Nutlets (styles) usually 2 or 3 (rarely l) 27. 

19. Inflorescence villous -hairy; shoots and leaves usually also rather 
densely pubescent 20. 

+ Inflorescences glabrous; flowers sometimes with hairy lower part 

of hypanthium; leaves glabrous or subglabrous 21. 

20. Leaves oblong-cuneate with narrowly cuneate decurrent base; 

lobes at level of upper third of blade 

17. C. sphaenophylla A. Pojark. 

+ Leaves broadly obovate or broadly rhomboid in outline; lower 

lobes at middle of blade 39. C. armena A. Pojark. 

21. Flowers in compound corymbiform inflorescences 23. 

+ Flowers in simple umbellate inflorescences (rarely single pedicels 

2- or 3 -flowered) 22. 



See also C. ti ansch anic a A. Pojark.; fruits small, 6 — 10mm in diameter; leaves 1.5— 3 cm or 3 — 5cm 
long, ovate, incised to l / z — % or else deeply partite with acuminate, unequally serrate lobes. This note 
also pertains to C.altaica Lge. 



319 



22. Fruiting sepals spreading; styles (nutlets) 2 or 3; nutlets with deep 
sinuate lateral groove; leaves with obtuse, broad, little -developed 
lobes in upper part of blade * C . oxyacantha L. 

+ Fruiting sepals erect; style (nutlet) 1; nutlet nearly smooth 

laterally; leaf lobes well developed, acute or obtuse 

31. C. microphylla C. Koch. 

23. Leaves slightly lighter below than above, without waxy bloom below, 
and with very prominent network of veins above; leaf lobes usually 
serrate from the middle; sepals narrow, long-acuminate; fruits 
oblong -ellipsoid; styles usually bent in upper part. (Series 
Kyrtostylae) 26. 

+ Leaves conspicuously bicolor, whitish with a waxy bloom below; 

veins not prominent; leaf lobes usually dentate only in upper part or 
else entire; sepals broader, more shortly acuminate, often subobtuse; 
fruits broadly ellipsoid to subglobose; style erect. (Series Mono- 
gynae) 24. 

24. Leaves dark green (olive-green) above, very lustrous; upper leaves 
on fruiting shoots mostly 5 -partite, the lobes directed forward and 
usually approximate in upper part 28. C. monogyna Jacq. 

+ Leaves light, bright green or gray-green, usually tripartite, the lower 
lobes separated by broad notches, ascending at an angle of 45° ... 25. 

25. Fruits 9 — 11mm long; upper leaves on fruiting shoots 2.2 — 3.5cm 
long, mostly tripartite, the middle lobe often tripartite at the apex, 
usually conspicuously tapering toward base, its length 1.5 — 2 times 
as long as or rarely equaling its width; notches at middle of blade 

or lower, rarely higher; sometimes leaves 5 -partite 

29. C. pseud oheterophy 11a A. Pojark. 

+ Fruits 12 — 13 mm long; upper leaves on fruiting shoots 3 — 5 cm long, 
tripartite, the middle lobe usually not conspicuously tapering toward 
base, broader, its length equaling, rarely to 1.5 times greater than 

its width; notches conspicuously higher than middle of blade 

30. C. turcomanica A. Pojark. 

26. Upper leaves on fruiting shoots to 5.5 cm long, 5 — 7-partite; lower 
part of hypanthium glabrous 24. C. kyrtostyla Fingerh. 

.„„ + Upper leaves on fruiting shoots smaller, to 3.5 cm long, 5-partite; 

lower part of hypanthium usually hairy 

2 5. C. turkestanica A. Pojark. 

27. Fruits yellow, with 2 or 3 nutlets; inflorescences and young shoots 
tomentose-pubescent 12. C. pontica C. Koch. 

+ Fruits red or black 28. 

28. Inflorescences glabrous, only lower part of hypanthium sometimes 
hairy 32. 

+ Inflorescences densely villous-hairy to tomentose 29. 

29. Upper leaves on flowering shoots deeply (to % — %) 5 — 7-partite, the 
lobes oblong, with few large teeth only near the apex or entire; 
notches between lobes narrow, the lowest at level of lower third of 
leaf; leaf pubescence usually dense 13. C. meyeri A. Pojark. 

+ Upper leaves on flowering shoots usually dissected to % — 3 U into 3 — 5 
lobes or else 7-fid to l lz~~ X l2'> lobes and notches broader, the notches 
arranged not lower than middle of leaf 30. 



320 



30. Upper leaves on flowering shoots dissected to / 3 — / 2 into 5 — 7 
lobes, the latter broad, with several teeth near the apex or entire . . 
14. C. eriantha A. Pojark. 

+ Upper leaves on flowering shoots .3 — 5-partite or lobed, the lobes at 

middle of blade . 31. 

31. Lobes serrate from the middle or only near apex, obtuse or acute; 

inflorescence densely whitish- villous -tomentose 

15. C taurica A. Pojark. 

+ Lobes usually serrate nearly from base, acute or acuminate; 

inflorescence villous -hairy but not tomentose 

16. C. ucrainica A. Pojark. 

32. Fruits small, 8 — 10 mm long, oblong, black-purple, with 1—3 nutlets; 

leaves to 3.5 cm long, pinnately 5-partite 

35. C. zangezura A. Pojark. 

+ Fruits 11— 18 mm long, subglobose 33. 

33. Leaves 5 — 7-lobed, with broad obtuse lobes, the upper leaves little 

developed, with few large teeth only near the apex 

20. C. transcaspica A. Pojark. 

+ Leaves partite with acute lobes 34. 

34. Leaf lobes with few large teeth only near the apex 3 5. 

+ Leaf lobes unequally serrate from the middle or nearly from 

base 37. 

35. Leaves dark green (olive-green) and lustrous above, much lighter 
below 34. C. dipyrena A. Pojark. 

+ Leaves light glaucous -green, only slightly lighter below 36. 

36. Fruits purple-black; shoots spiny 18.C. ambigua C. A. M. 

+ Fruits dark blood-red; shoots unarmed 

22. C. atrosanguinea A. Pojark. 

37. Fruits dark red; upper leaves on flowering shoots 7 — 9-lobed, 

the lobes serrate, not incised 19. C. volgensis A. Pojark. 

+ Fruits blackish purple or purple-black 3 8. 

3 8. Spiny shrubs; leaves broad, with rounded base, the lower lobes 

twice as long as the upper, all lobes unequally serrate, rarely with 

deep incisions 21. C. caucasica C.Koch. 

+ Unarmed trees; leaves more deeply dissected, the lobes coarsely, 

unequally, and acutely dentate, often — mainly the lower — with deep 

incisions 39. 

39. Leaves large, to 7 cm long on flowering shoots; fruits large, 

12 — 15mm long, subglobose 23. C. songorica C.Koch. 

+ Leaves much smaller; fruits more oblong and smaller 

38. C. pseudo-ambigua A. Pojark. 

Section 1. PINNATIFIDAE Zbl. in Beissn., Schelle u. Zab., Handb. d. 
Laubholzben. (1903) 178; C K. Schn. 111. Handb. d. Laubh. I (1906) 769.- 
Fruits with 3 — 5 nutlets, smooth laterally, large, with compact (not mealy) 
flesh, red with white warts. 

1. C. pinnatifida Bge. in Mem. Acad. St. Petersb. II (1831) 100; Kom., 
Fl. Mansh. II (1904) 466; Kom. and Alis., Opred. rast. Dal'nevost. kraya II 
(1932) 637.— Mespilus pinnatifida C. Koch, Dendrol. I (l 869) 1 52. — 
C. oxyacantha var. p inn at i f i d a Rgl. in A. H. P. I (1871—1872) 118.- 
Ic: Kom. and Alis., 1. c, Plate 193. 



321 



422 



Tree or tall branching shrub to 6 m high; spines few, 1—2 cm long, 
often absent; young shoots glabrous or hairy; new annotinous shoots light 
brown, the bark of older branches dark gray; leaves bright green, lustrous, 
glabrous or hairy along the veins below, with beards in angles of main 
veins, cuneate or truncate at base, ovate or oblong-ovate, to 6 — 8.5cm long 
and 5 — 6.5 cm broad on flowering shoots, larger on sterile shoots, deeply 
pinnatifid into usually 3, less often 4 or 2 lobes on each side; lobes acute 
or acuminate, oblong -triangular, serrate; inflorescences pendulous, 
12 —20 -flowered, the axes and pedicels glabrous (v. g e ho 1 e n s i s C. K. 
Schn.) or with more or less dense to tomentose pubescence (v. p i 1 o s a 
C. K. Schn.); flowers 8 — 12 mm in diameter, with white petals turning 
pink toward end of anthesis; sepals sharp-pointed, reflexed at anthesis and 
in fruit; styles 3 — 5; fruits lustrous, bright red with light warts, to 1 7 mm 
long, obovate or rounded-ovate. Fl. May, fr. from end of August. (Plate 
XXVIII, Figure l). 

Riverbanks, on sandy soils, stony slopes; solitary trees or in tree and 
shrub thickets. — Far East: Ze. -Bu., Uss. Gen. distr. : Jap. -Ch., Manchuria, 
Korea, N. China. Described from Chihli Province. Type in Leningrad. 

Economic importance. Cultivated in parks in the European part of the 
Soviet Union, but seldom, although this is one of the most ornamental 
representatives of this genus. 



Section 2. SANGUINEAE Zbl. in Beissn., Schelle u. Zab., Handb. d. 
Laubholzben. (1903) 174 (ex parte). — Eusanguineae Rehder in Frut. 
Vilm. Cat. prim. (1904 — 1905) 111.— Section Sanguineae C.K. Schn., Ill 
Handb. Laubh. 1 (1906) 77. Fruits mealy, with 3 — 5 nutlets, strongly 
compressed and pitted laterally. 



Cycle Sanguineae A. Pojark. — Fruits red or yellow, with yellowish, 
mealy flesh; inflorescences glabrous. 

2. C. sanguinea Pall. Fl. Ross. I, 1 (1784) 25; Ldb., Fl. Ross. II, 1, 88 
(ex parte: excl. pi. dahur. ); Shmal'g., Fl. I, 3 50; Voronov in Fl. Yugo- 
Vost. V (1931) 496; Kryl., Fl. Zap. Sib. VII, 1466. — C r a t a e gu s sangui- 
nea var. ty pic a Rgl. in A. H. P. I (1871—1872) 115 (excl. syn. plur.).— 
Mespilus purpurea Poir. in Lam. Encycl. Suppl. IV (1816) 73. — 
M. sanguinea Spach, Hist. veg. II (1834) 62. — Ic: Pall., 1. c, tab. XI. 

Tall shrub, less often small tree, 1 —4m high; shoots strong, purple- 
brown, lustrous, usually with thick, straight spines 2.5—4 cm long; stipules 
falcate or obliquely cordate, coarsely glandular-dentate; leaves on short 
shoots 2 — 6 cm long, 2.5 — 5 cm broad, those on vegetative shoots to 9.7 cm 
long, usually hairy on both sides, dark green above, much lighter below, 
obovate to broad-rhomboid, acute, usually cuneate and entire at base, 
shallowly 3 — 7-lobed or coarsely dentate; lobes serrate; leaves on sterile 
shoots sometimes more deeply lobed or parted, occasionally almost 
dissected at base; inflorescences rather dense, 3— 4 cm long, 4 — 5 cm in 
diameter, with caducous filiform bracts; pedicels and outside of hypanthium 
often slightly hairy; sepals oblong -triangular, entire or with 1 or 2 teeth; 
flowers 12 — 15 mm in diameter; stamens 2 0, with purple anthers; styles 



322 



usually 3 less often 4, exceptionally 2 or 5; fruits 8 — 10 mm in diameter, 
blood-red, very rarely orange-yellow (v.chlorocarpa C. Koch), pellucid, 
with 3 or 4 nutlets and with mealy flesh. Fl. May— June, fr.from August. 
(Plate XXVIII, Figure 4). 

Open forests, forest edges, riverbanks and riverain deciduous forest 
strips, in the forest and forest- steppe zones and at the edges of the steppe 
zone.— European part:U. V.— E.part, in the former Aleksandrov and 
Pereyaslav counties (Flerov) — W. border; V. -Kama — W part as far as 
Glazov, V. -Don, Transv. ; W.Siberia: to 62°N ; Ob Irt., Alt (except E part); 
E. Siberia: S.half), U. Tob., Yenis. (S part), Lena-Kol. (S.part), Ang. -Say., 
Dau. (W. part, Selenga and Barguzin); Centr. Asia: DzurTarb. and Balkh . 
Gen.distr.: Mong.-Tuva ASSR and N. Mong. (as far east as Urga [Ulan 
Bator]). Described from the S.Urals. Type in London. 

Note. In regions where the distribution areas of C.sanguinea 
coincide with those of C.altaica Lge. and C.dahurica Koehne, these 
species form hybrids intermediate in their characters between the parental 
species. 

Economic importance. Most widespread species of hawthorn cultivated 
in the Soviet Union; mainly used for hedgerows. 

3. C.dahurica Koehne ex C. K. Schn., Handb. d. Laubh. I (1906) 773; 
Fedde, Repert. Ill (1907) 224; Kom . and Alis., Opred. rast. Dal'nevost . kraya 
(1932) 637.- C. sanguinea (non Pall.) Turcz., Fl. baic. -dah.I, 408.- 
C.sanguinea a genuina et6 glabra Maxim., Prim. fl. amur. (1859) 
101; Kom., Fl. Manch.II (1904) 469.- C. chitaensis Sarg., PI. Wilson. I 

2 (1912) 183.- Ic: C. K. Schn., 1. c ., f. 457, n-o. 

Tree or small shrub 2 — 6m high; shoots dark purple, unarmed or with 
spines 1—2.5 cm long; leaves mostly glabrous on both sides, rarely with 
short coarse pubescence above or even on both sides, those on short shoots 
1.8 — 4.5 cm long, 1 .2 — 3 cm broad, oblong-obovate or oblong- rhomboid, 
cuneately decurrent along petioles, with (5) 7 — 9 large (sometimes little 
developed) teeth or small lobes on each side, serrate; leaves on sterile 
shoots much larger and more deeply lobed, sometimes nearly partite with 
truncate base; inflorescence 7 — 20-flowered; bracts filiform, caducous; 
sepals oblanceolate, entire or with 1 or 2 teeth on each side, reflexed, 
shorter than hypanthium; flowers ca. 1.5 cm in diameter, with purple anthers 
(stamens 20); styles 2 — 4; fruits small, 6 — 8 mm in diameter, globose or 
slightly oblong, red or orange-red, apparently occasionally orange-yellow, 
with 2-4 nutlets pitted ventrally. Fl. May, fr. August. (Plate XXVIII, 
Figure 2). 

Deciduous forests, forest edges, dry mountain slopes among trees and 
shrubs, river valleys, flooded meadows. — E. Siberia: Yenis. (Bunyukonskii 
and Onaon ranges), Lena-Kol.; nearly to 65° N., Ang. -Say . (E. part); Dau.; 
Far East: Uda, Ze. -Bu., Uss. (Usually on the Amur, very rarely in S. part 
of region). Gen.distr.: Jap. -Ch. (Manchuria), Mong. (N. Mong., Urga, 
Kentei Range). Described after a cultivated specimen. Type in Berlin. 

Economic importance. Occasionally in arbor etae; not widely cultivated. 

4. C.altaica Lge., Revis. spec. gen. Grataegi (1897) 42 (excl.var. 
villosa); C.K. Schn., Illustr. Handb. Laubh. I (1906) 773; Fedch., Consp. 
fl.turk.III (1909) 41; Kryl., Fl. Zap. Sib. VII, 1467.- C.sanguinea var. 



323 



[425) 




PLATE XXVIII. 1— Crataegus pinna t if id a Bge.: a - b) nutlet; 2-C.dah uri ca Koehne: 
a— b) nutlet; 3— C.altaica Lge.: al flower, b) fruit.c— d) nutlet, e) leaf of C.altaica 
var. incisa Rgl., Pall., leaf; 5-C.chlorosarca Maxim. : a - bl nutlet. 



324 



inermis Kar. et Kir. in Bull. Soc. nat. Mosc. XIV (1841) 328; XV (1842) 351; 
Ldb., Fl.Ross. II, 1, 88.- C-pur pur e a 2.altaica Loud., Arb. Brit. II 
(1844) 825.— C.sanguinea var. xanthocarpa et var. inc i s a Rgl. in 
A. H. P. I (1871-1872) 116.- C.songorica Rgl. in Delect. Sem. Hort. 
Petr. (1876) 36 (nom.nud.), non C. Koch. — C.sanguinea var. songorica 
Rgl. in sched. — C.pinnatifida Franch. (non Bge.) in Ann. Sc. Nat. 6 
ser., XVI (1883) 288.-C.korolkovii L. Henry in Rev. hort., Nouv. ser. 
LXXIII (1901) 308; M.Popov in Tr . Prikl. Bot., Sel.i Gen. XXII, 3 (1929) 
436.— C.altaica var . h i s s a r i c a Bornm . in sched.— Ic: Lge., I.e., 
tab. V; L.Henry, 1. c, tab. col. A - F 1 ; C. K. Schn.,l.c, fig. 437 i, 438 k-m. 

Small tree, usually unarmed, rarely with few thick 0.6 — 2 cm straight 
spines; annotinous shoots glabrous, lustrous, brownish red, beset with light 
lenticels; older shoots yellowish gray or reddish gray; stipules large, 
cordate or falcate, with large teeth terminating in a gland; leaves usually 
broadly triangular-oval to orbicular (f.latifolia M. Pop. in sched.), from 
3.5 cm (at base of flowering shoots) to 12 cm long, 2.5 — 10 cm broad, on the 
average 6.5 cm; leaves acute, broadly truncate, less often rounded, shallowly 
cordate, or rounded -cuneate at base, dull, glaucous-green above, lighter 
below, mostly subglabrous, less often sparsely puberulent above, shallowly 
7 — 9-lobed, the lower horizontal lobes usually much larger than the others, 
sometimes subentire; rarely but throughout the distribution area there are 
specimens with leaves deeply lobed (especially on sterile shoots), nearly 
pinnatisect at base, sometimes even with remote lower lobes: C.altaica 
v.incisa C. K. Schn. (C. s a n g u i n e a v.incisa Rgl., C. w a 1 1 i an a v. 
incisa C.K.Schn.); leaf margin acutely dentate; inflorescences 20 — 50- 
flowered, glabrous; pedicels ca. 6.5mm long;sepals triangular-lanceolate, 
curved, shorter than and appressed to the hypanthium, entire or with 1 or 
2 small teeth; flowers white; stamens 20; anthers white; styles nearly 
always 5 (very rarely 4); ovary apex sparsely hairy. Fruiting abundantly, 
slightly nodding; fruits 8 — 12 mm in diameter, globose or slightly 
flattened above, rarely oblong, their color varying at maturity from ocher- 
or orange-yellow (f. fl av a M. Pop.) to orange-brown (f. fu s c a Lge., 
f.rubescens M. Pop.), with very soft, mealy, light yellow, tasty flesh. 
Fl.May- June, fr. August- September. (Plate XXVIII, Figure 3). 

Forests on mountain slopes and ravines, in the middle mountain zone.— 
European part: L. V., vicinity of Ural'sk in the chalky mountains: an 
insular habitat, the northwesternmost limit of the species: W.Siberia: Alt. 
(village of Ul'binskii); Centr.Asia: Ar. -Casp. (Mugodzhary to Ulu-Tau), 
Dzu. -Tarb., Pam . -Al. : N. slope of the Alai Range, Shugnan, Darvaz, Gissar, 
Zeravshan; throughout Tien Shan (very frequent in the W., rare in Centr.) 
Gen.distr.: apparently Afghanistan. Described after a cultivated specimen 
reputedly from the Altai. Type unknown. 

Economic importance. Rather common in cultivation but usually 
confused with C. s anguine a Pall. 

Note. A hawthorn very closely related to C.altaica has been 
described from [former] British Baluchistan as C.wattiana Hemsl.et 
Lace (Journ. Linn. Soc. XXVIII, 1891, 323, tab. 40). Schneider considers it 
identical to C.altaica Lge . (Fedde, Repert . Ill, 1906, 229), Rehder, however, 
(Man. Cult. Trees a. Shr . 763) distinguishes two independent species, 
indicating a number of differences. Owing to the lack of material from 
Baluchistan, it is better to adopt the second opinion. 



325 



Hybrids of C.altaica Lge.X C.pontica C. Koch have been collected 
in W. Tien Shan. The specimens examined have white-tomentose-pubescent 
young shoots, axial parts of inflorescence, and petioles; leaves pubescent 
below or on both sides, usually resembling in shape those of C.altaica, 
less often those of C.pontica; fruits (immature) small, with 5 pitted 
nutlets, slightly flattened at the poles. 

C.dshungarica Zbl. (p. 351) is a hybrid of C.altaica Lge. X 
C.songorica C.Koch. Hybrids of C.altaica Lge.X C.sanguinea 
Pall, are rather common in regions where the two distribution areas 
coincide. 

C.tianschanica A. Pojark. (p. 351) is probably a hybrid of 
C.altaica Lge.X C.turkestanica A. Pojark. 

5. C.maximowiczii C. K. Schn., Handb. d. Laubholzk. (1906) 771; Kom. 
~ in A. H. P. XXXIX, 1 (1923) 74; Kom . and Alis., Opred. rast . Dal'nevost . 

kraya II (1932) 637.— C.sanguinea v.villosa Maxim., Prim. fl. amur . 
(1859) 101; Kom., Fl.Mansh.II (1904) 469.- C.altaica v. villosa Lge., 
Rev. Sp. gen. Grat. (1897) 42.- Ic: C. K. Schn. 1. c, f. 437a - b; 438a-c. 

Shrub or small tree to 7 m high; young shoots with dense spreading hairs, 
annotinous shoots subglabrous, with lustrous reddish-brown bark; spines 
absent or few, 1 .5 — 3.5 cm long, thick; stipules large, lanate, cordate at 
base, incised-dentate; leaves on short shoots 3.5 — 8 cm long, 2.5 — 5 cm 
broad, obovate, with cuneate base, scattered-hairy above, more or less 
(sometimes very densely) velvety -pubescent above, shallowly 9— 13-lobed 
or incised, the lobes often obtuse; margins unequally serrate; leaves on 
vegetative shoots to 13 cm long, 10 cm broad, mostly deeply trilobate; 
inflorescences rather dense, corymbiform, to 5 cm in diameter, with densely 
hairy axes and pedicels; flowers ca. 1.5 cm in diameter; hypanthium and 
sepals tomentose; stamens 20; styles 3 — 5; fruits initially hairy, later 
glabrous, red. Fl.end of May, June, fr.from August. 

Riverrain deciduous forest strips, crests, meadows of flooded valleys, 
forest edges, dry mountain slopes; solitary trees or shrubs.— E.Siberia: 
rarely; Ang. -Say. (near Irkutsk), Dau. (on the Shilka River near Sretensk); 
Far East: Uda, Ze. -Bu., Uss., Sakh. Described from the Amur. Type in 
Leningrad . 

Economic importance. Rarely cultivated. 

6. C.remotilobata H.Raik.ex M. Pop. in Bull. Appl. Bot., Gen. a. Plant 
breed. XXII, 3(1929)438. 

Small tree; young shoots glabrous; annotinous shoots dark, reddish brown, 
lustrous, older shoots with mottled brownish-gray bark; spines slender, 
0.6 — 2.5 cm long; leaves (2.5) 3 — 7 cm long, firm, oblong-ovate, broadly 
truncate at base, gradually acuminate, glabrous on both sides, usually deeply 
7-partite, often dissected to base, the lower lobes sometimes quite separate, 
less often the blade incised into 7 lobes only to the middle; lobes oval- 
lanceolate to oblong-lanceolate, acuminate, usually acutely denticulate only 
on outer margin, less often only slightly crenate or even entire; petioles 
1.5 — 2.5 (3) cm long, glabrous; inflorescences 25 — 70-flowered, glabrous; 
sepals broadly triangular to triangular-lanceolate, acute, slightly shorter 
than hypanthium; stamens 20, with white anthers; styles 4 or 5; fruits red, 



326 



small, 4 — 6 mm, rarely 6—9 mm, globose, red; nutlets 3 — 5, triangular, 
strongly pitted laterally, 3 — 4 mm long. Fl. May, fr. from end of August. 

Riverbanks. — Centr. Asia: Pam. - Al. (Khodzhent, Kanibadam, and Gul'cha) 
and W. Tien Shan, in the Chatkal River valley; known mainly from the 
cultivated area of the Tashkent oasis, where it is often found in irrigation 
ditches and at roadsides. Endemic. Described from the vicinity of 
Khodzhent. Type in Leningrad. 

Note. C.remotilobata is clearly differentiated from C.altaica 
by its smaller red fruits with 3 or 4 very small nutlets; inflorescences with 
more branches and more flowers, and leaves from very deeply dissected to 
partite (resembling those of C. alt a i c a var.incisa but much smaller). 



Series Nigrae A. Pojark. — Fruits black with greenish flesh; nutlets 
4 or 5, with large hypostyle, longitudinally grooved on back and sides. 
Apart from the USSR species, this series includes C. nigra W. et K. — 
which was erroneously referred to the section Pentagynae — and 
C.jozana C. K. Schn. in Japan. 

7. C.chlorosarca Maxim, in Bull. Soc. Nat. Mosc. LIV, 1 (1879) 20; 
C.K.Schn.,Handb.Laubholzk.I (1906) 173; Kom., Fl. Kamch. II (1929) 
235; Hulten, Fl. Kamtch. Ill (1929) 49.- C.oxyacantha fructu rubro 
et nig r o, Krashenn., Kamch. 2nd edit. (1786) 309.— C.oxyacantha 
(non L.) Pennat, Arch. Zool. (1784) CXV; Georgi, Beschreib. Ill, 4, 1009 
(ex parte, quoad pi. kamtsch.). — C.sanguinea Pall, (ex parte, quoad, pi. 
kamtsch.) I (1784) 25; C. K. Schn., I.e., (ex parte, quoad pi. kamtsch. ). — 
C. m an d s hu r i c a hort.Buek. Vernacular name: kharem. 

Tree to 6 m high, with dense crown; branches erect, flexible, frequently 
knotty; bark on young shoots dark purple, on old shoots gray; spines short, 
1 — 1.5cm; stipules falcate, broad, dentate; leaves 4.5 — 8 cm long, to 
3.5— 8 (9) cm broad on short shoots, to 13 cm long and 10 cm broad on long 
shoots, glabrous on both sides or scattered-hairy above and rather densely 
pubescent below, or else hairy on both sides, oval, broadly cune ate, less 
often* truncate at base, shallow ly 9 — 11-lobed or incised, serrate; 
inflorescences 4 — 9 cm in diameter; 1.5 — 3 cm long, few-flowered, glabrous, 
much shorter than leaves; sepals lanceolate-triangular, dentate, 
taper-tipped, half as long as or equal to hypanthium, glabrous or hairy 
inside; petals orbicular, clawed; stamens 20; styles 5; fruits black at 
maturity with greenish flesh, red when immature, with 4 or 5 nutlets pitted 
at the sides. Fl. July, fr. August- September. (Plate XXVIII, Figure 5). 

Dry river terraces, edges of riparian forests, only in the forest zone 
and not very close to the sea; solitary or in groups of 2 or 3 trees.— Far 
East: Sakh., Kamch. (except N.). Gen.distr.: Jap. (Hokkaido). Described 
after a cultivated specimen. Type in Leningrad. 

Note. Maximowicz, quoting Krasheninnikov, cites in his list of 
Kamchatka plants a red- fruited as well as a black- fruited hawthorn. He 
believes that the latter may be C.sanguinea v. glabra Maxim.; this, 
in fact, is a synonym of C.dahurica Koehne, which does not grow in 
Kamchatka . It is likely that specimens of C.chlorosarca with 
immature fruits were mistaken for red- fruited hawthorn; a less likely 
possibility is that he had in mind Sorbus sambucifolia. 



327 



Economic importance. Ornamental, rarely cultivated in the Soviet Union; 
its substitute species .C.nigra C.etK.- distinguished by the dense 
velvety pubescence of underside of leaves and inflorescence, different leaf 
shape, and smoother nutlets - is more widely cultivated. 

Section 3. PENTAGYTSTAE Zbl. in Beissn., Schelle u. Zab., Handb. d. 
Laubholzben. (1903) 178; C. K. Schn., 111. Handb. d. Laubh. I (1906) 277.- 
Fruits black, with little- developed reddish flesh, the 3 — 5 triangular nutlets 
smooth on the sides and with inconspicuous longitudinal grows on dorsal 
side. Two species: C . p e nt ag yn a W. et K. and C. p s e ud o me 1 an o - 
carpa M. Pop. 

8. C.pentagyna W. et K.in Willd., Sp. pi. II, 2 (1799) 1006; Rgl.in 
A.H.P I (1872) 113; Grossg., Fl. Kavk. Iv (1934) 290. — C . melanocarpa 
M.B.,Fl.taur. -cauc. I, 386; Ldb., Fl. Ross. II, 1, 89; Boiss., Fl.Or.II (1872) 
661 (excl. var.heterophylla); Medv., Der. i kust. Kavk. (1919) 1 10. - 
C.atrofusca Stev. ex Hoh., Enum. pi. Talysch (183 6) 130 (nom.nud.).-?. 
C.oliveriana Bosc in DC., Prodr. II (1825) 63 0.— C.oxyacantha var. 
oliveriana Lindl. in Bot. Reg. XXIII (1837) sub tab. 1933.- M e s p i 1 u s 
pentagyna Spreng., Syst. II (1825) 507; Shmal'g., Fl.I, 350— C. c olc h i c a 
Grossh., Fl. cauc. IV (1934) 290.- Ic: Rchb., Ic. Fl. Germ. XXV, tab. 101. 

Tree 3 — 8 (l2)m high, less often a tall shrub; young shoots lanate- 
pubescent, less often glabrous; bark of branches gray; spines thin, 5 — 10 mm 
long, usually few; leaves dark green, lustrous, and hairy above, lighter and 
dull below, more or less pubescent, often almost to velvety- tomentose 
below (C.colchica Grossh.) or glabrous from the very beginning (v. 
glabrata Trautv., var. at r o f u s c a Boiss.); broadly cuneate or truncate, 
sometimes slightly notched at base; leaves on flowering shoots 2 — 5.5 cm 
long, 1.5 — 5 cm broad, the lower usually trilobate, the others 5 — 7-pinnatifid; 
lobes obtuse or pointed, acutely dentate near the apex; leaves on sterile 
shoots larger and often broader, to 8 — 9 cm long and broad, more deeply 
dissected, the lower lobes sometimes incised; inflorescences to 10 cm in 
diameter, some with repeatedly branching axes of the second order, many- 
flowered; pedicels, hypanthium, and outside of sepals usually more oi*less 
hairy, sometimes even tomentose; sepals ovate- triangular or broadly 
4 oi triangular, mostly with a short sharp point, less often acute, spreading at 
anthesis, erect in fruit; corolla 13 — 17 mm in diameter; stamens 20; styles 
3 — 5, quite free or more or less fused; fruits black with glaucous bloom, 
globose, the reddish flesh little developed; nutlets 3 — 5, smooth, triquetrous. 
Fl. May — June, fr. end of August — September. 

Forests and forest edges, among shrubs, in the Caucasus in the middle 
forest zone. — European part: several habitats in the Ukraine — M. Dnp. 
(Mgar near Lubny and Grebenniki), V. -Don (vicinity of Kharkov, Svyatye 
Gory and Slavyansk), Crim.; Caucasus: all districts (absent in Daralagez 
in Armenia). Gen. distr.: Centr. Eur. from Slavonia, Serbia, Bulgaria, and 
Banat to Podolia and the Balkan Peninsula (to Thrace and Macedonia), 
Turkish Armenia. Described from Hungary. Type in Berlin. 



328 



Note. Extremely polymorphic species with wide, discontinuous 
distribution area. Geographical separation cannot be ascertained on the 
basis of herbarium material; this question can be solved only by studying 
the variability of C. p ent a gyn a in the field in the various regions where 
it grows. C.pentagyna W.etK. apparently readily forms hybrids with 
the most varied species; hybrids of C. p e nt agyn a W. et K. XC.san- 
guinea Pall. = C. lamb e r ti ana Lge.,C.pentagyna W.etK. X 
C.crus galli L. =C.hiemalis Lge. Hybrids of C.pentagyna 
usually have dark fruits and can be recognized by the reddish color of the 
fruit flesh, by the deep umbilicus, and by the ascending fruiting sepals. 
The following hybrids of C.pentagyna are often found in the Soviet Union: 

C.pentagyna W.etK.XC.kyrtostyla Fingerh. - These hybrids are 
most often found in the Crimea, N. Caucasus, and W. and S. Transc. (Zangezur), 
E. Transc. (districts of Kirovabad and Kuba), and Talysh. The hybrids are 
mostly intermediate: fruits usuaUy ellipsoid, from dark red to purple- black 
(without bloom), the flesh usually colored, with 2 or 3 nutlets (with slight 
admixture of fruits with 1 or 4 nutlets); in leaf shape, pubescence, shape and 
position of sepals they show a range of transitions between the two species. 
These hybrids are usually cited as C.oxyacantha. 

C.pentagyna W.etK.XC.monogyna Jacq., collected in the Crimea 
and the N.Caucasus. These hybrids have dark red fruits, with mostly 2, 
rarely 1 or 3 nutlets; leaves bicolor, often with waxy bloom below, with 
few-toothed, mostly subobtuse lobes as in C. mono gyn a. Quite frequent 
in the Crimea; according to available material they show great consistency 
of characters and, therefore, should be described under a separate name — 
C.dipyrena A. Pojark. (p. 351). Pallas's C. d i g yn a Pall. (Ind. taur. 
1796, p. 107 — nom. nud.) may belong here. 

C.zangezura A. Pojark. (p. 352) is one the hybrids whose parental 
species is C.pentagyna W.etK. 

C.pentagyna W. et K.XC.orientalis Pall. — I consider as a hybrid 
of these species C. s c hr ad e r iana Ldb. (p. 353), described (by Schrader 
as S.sanguinea Schrad. and renamed by Ledebour) after cultivated 
specimen grown from seeds obtained from E.Crimea. Hawthorn identical 
to the Schrader specimen has been discovered in Zangezur, where it 
shows consistency of characters, being, apparently, an hereditarily stable 
form. 

9. C.pseudomelanocarpa M. Pop. sp.n. in Addenda VIII, p. 384, Linchevskii, 
Rast. Zap.Kopet-daga, Rast. resursy Turkm. I (1935) 59 (nom.mud. ).- 
C.melanocarpa M. Pop. in Bull. Appl. Bot., Gen. Plant, breed. XXII, 3 
(1929) 436 (non M.B.). 

Small tree with densely pubescent, almost tomentose shoots and branches 
covered with light gray bark; spines 5 — 20 mm long, straight; leaves light 
green, 14 — 35 mm long, 8 — 35 mm broad, those at base of short shoots 
cuneate or oval-cuneate, shallowly trilobate or trifid only at the apex, the 
others 5 — 7-lobed, oval, mostly truncate or rounded-truncate at base, less 
often rounded- cuneate; leaves on sterile shoots more deeply parted, 



329 



433 



sometimes almost dissected at base; lobes directed forward, horizontal 
only on sterile shoots, entire or with few teeth above the middle; 
inflorescence dense, to 6 cm in diameter, the axis more or less lanate- 
pubescent; pedicels and hypanthium subglabrous; ovary mostly slightly 
hairy; flowers 9 — 12 mm in diameter; stamens 20; styles 5; sepals 
recurved in young fruits; mature fruits black, without bloom, 7 — 8 mm in 
diameter. Fl. June, fr. September. 

Ravines. — Centr. Asia: Mtn. Turkm. (Kara-Kala District). — Gen. distr.: 
Iran, -mountains of the S. shore of the Caspian (Asterabad and Mazanderan). 
Described from the Ioldere Gorge. Type in Leningrad. 

Note. Distinguished from C.pentagyna by differently shaped, 
smaller, lighter leaves, smaller flowers, fruits without glaucous bloom, 
and larger spines. 

C.pseudomelanocarpa M. Pop., like its substitute species 
C.pentagyna W. et K. produces hybrids with species of other sections 
growing in the same sites . C.pseudoazarolus M. Pop . (p . 3 54) is a 
hybrid of C.pseudomelanocarpa XC.pontica C. Koch. There are 
apparently hybrids of C.pseudomelanocarpa XC.turcomanica; 
the specimens from W. Kopet Dagh collected in the Ioldere Gorge 
apparently also belong here: flowers 1 — 3, pistillate, leaves cuneate, 
glaucous below, with lobes closely approximate in upper part of blade. 

C. p s eudoambigua A. Pojark. (p. 355) with black fruits and 2 nutlets 
is a hybrid of C.pseudomelanocarpa M.Pop. XC.turkestanica 
A. Pojark. 

Section 4. AZAROLI Loud., Arb. et frut. brit., II, ed. 2 (1844) 326. - 
Orientales Zbl. in Beissner, Schelle u. Zabel., Hand. d. Laubholz. 
(1903)179; C. K. Schn., Illustr. Handb. d. Laubholz. I (1906)781. - 
Inflorescences tomentose-pubescent, compact, with short axes and pedicels; 
fruits yellow or reddish orange or (?) red; nutlets 2—5, smooth ventrally 
or laterally; anthers white; petioles (V 10 ) Vs - 73 as long as the blade. 

Economic importance. The fruits of species of this section deserve 
more attention than those of other sections of this genus, since they are not 
only larger and more fleshy but also have a pleasant taste. 



Series Orientales A. Pojark.— Fruits reddish orange, with 5 (4) nutlets; 
sepals recurved. 

10. C. orientalis Pall. exM.B, Fl. taur.-cauc. 1(1808)387; 111,322; 
Pallas, Index taur. (1796) 107 (nom. nudum); Grossg., Fl. Kavk. IV (1934) 
290.— Mespilus odoratissima Andr. Bot. Repos. IX (l 81 0) tab. 
590.- C. tanacetifolia taurica DC, Prodr. II (1825) 629.- 
C . tanac etif olia Ldb., Fl. Ross. II, 1, 90 (non Pers.); Steven in 
Bull. Soc. Nat. Mosc. XXIX, 1 (1856) 248.- Phaenopyrum odora- 
tissimum Roem., Syn. monogr. (1847) 158. — C. . t ana c e t i f o 1 i a var. 
orientalis Rgl. in A. H. P. l(l87l)ll4; Shmal'g., Fl. I, 350.- Ic: 



330 



Bot. Reg. (1836) tab. 1885; Bot. Mag. tab. 2314; Vol'f and Palib., Der. I 
kust., figure on p. 449 (leaves).— Vernacular names: gevizh' (Armenian, 
vicinity of Erevan), tsiteli gambro (Georgian, Ateni). 

Branching shrub 1— 2.5(3 )m high or small tree 3—5 m high, very rarely 
larger, with densely pubescent-tomentose shoots, of which many transformed 
into leafy spines; leaves firm, dull, with dense soft grayish pubescence on 
both sides, 3— 8(10) times as long as the petiole, oblong-ovate or obovate, 
cuneate at base; lower leaves on flowering shoots trilobate, the others 
5— 7 -fid, often very deeply so, nearly to midrib; lobes usually narrow, 
usually 3—3.5 times as long as broad (2.5— 4.1), with 1—4 teeth at the apex; 
lower lobes sometimes with a deep lobelike incision; leaves on sterile 
shoots broader, often with truncate base; inflorescence very compact, 
few-flowered, densely whitish-tomentose, with very short axes and pedicels 
2— 3(5) mm long; sepals lanceolate-triangular, long-acuminate, sometimes 
tapering to a subulate sharp point; corolla 15— 20 mm in diameter; 
stamens 20; styles 5, rarely 4; fruits 13— 20 mm in diameter, strongly 
434 flattened at the poles, pentagonal, reddish orange, slightly hairy or glabrous, 
with mostly 5, rarely 4 triquetrous nutlets. Fl. June, beginning of July, 
fr. from September. (Plate XXIX, Figure l). 

Middle mountain zone, dry stony slopes covered with shrubs.— European 
part: Crim. — S. and N. slopes of the Crimean Mountains, as far east as 
Feodosiya; Caucasus: S., S., and W. Transc. (Kura River valley and further 
south), Tal. Gen. distr.: Bal.-As. Min. and S. Greece. Described from 
the Crimea. Type in London. 

Economic importance. In the Caucasus the local population collects the 
fruits, which are distinguished by a pleasant, rather acid taste; they are 
found in markets and mostly eaten raw, less Often they are ground and 
mixed with flour for the preparation of a sweetish bread. Owing to its 
drought-resistance, this shrub may be of as great interest for arid regions 
as C. pontica C. Koch, since individual specimens yield fruits as large 
and fleshy as those of the latter species. These very dense and spiny shrubs 
can be recommended for hedgerows. The wood is used for tool handles. 

Series Szovitsianae A. Pojark. — Fruits (apparently red) with 3 or 4 
styles; fruiting sepals erect-spreading. One species. 

11. C. szovitskii A. Pojark . sp. n. in Addenda VIII, p. 385. — C . o ri en - 
talis 6 connecta Diapulis in Fedde, Repert. sp. no. XXXIV (1933) 
56 (ex parte). 

Shrub or small tree with brown-gray branches and crowded, very thick 
strong shoots densely tomentose when young, most shoots becoming 
shortish leafy spines; leaves firm, thick, glaucous, with almost continuously 
appressed-pubescent below, more sparsely so above, with petioles / 8 — / 6 as 
long as the blade; lower leaves on flowering shoots obovate -cuneate, 
coarsely dentate or incised only at the apex, leaves higher up trilobate but 
the upper leaves usually deeply (to % of blade) 5 -partite, broadly obovate - 
rhomboid; middle lobe often cuneately tapering toward base, acute or 



331 



obtuse, the lateral lobes with parallel margins, acute, rather broad (1.5—2.5: 1; 
rarely 3: l), with few teeth near the apex, less often entire; leaves on sterile 
shoots larger, deeply 5— 9-partite with narrower, more incised lobes; 
inflorescences to 5 cm in diameter, compact, 10— 12 -flowered, densely white- 
tomentose; pedicels 1.5— 5 mm long; sepals broadly triangular, with a 
435 strong cusp, erect at anthesis, erect-spreading in fruit; corolla ca. 18 mm 
in diameter; stamens 20; styles 3 or 4; fruits 12— 15 mm in diameter, 
slightly hairy; mature herbarium specimens dark red with usually 3 or 4, 
rarely 2 obtusely triangular nutlets; hypostyle triangular, attaining middle 
of nutlet. Fl. July, fr. October. 

Stony slopes among shrubs, forest edges in the middle mountain zone.— 
Caucasus: E. Transc. (in E. Karabakh: vicinity of Shusha, Dzhebrail, and 
Gadrut). Gen. distr. : Turkish Armenia (Kharput) and As. Min. (Tokat). 
Described from the vicinity of Shusha. Type in Leningrad. 

Note. Most closely related to C.orientalis Pall., from which it 
is distinguished by number of styles, by shape and arrangement of fruiting 
sepals, and by shape of broader-lobed leaves. 

Economic importance. Very ornamental, suitable for hedgerows, 
probably valuable as a fruit-bearing tree. 



Series Ponticae A. Pojark. — Fruits yellow, with 2 or 3 nutlets. 

12. C.ponticaC. Koch, Crat. et Mesp. (1854)49; Grossg., Fl. Kavk.; 
IV (1934) 290.- C. azarolus Fedtsch., Consp. Fl. turk. I (1909) 18 
(non L.); M. Popov in Tr. Prikl. Bot., Gen. i Sel. XXII, 3(1929)440; 
auct. plur. turkest. — Ic: M. Popov, 1. c, Fig. 98.— Vernacular names: 
dulyana (Tadzhik), alyuch (Turcoman), tetri-gambro (Georgian, in Ateni); 
dulyana, dulena (Uzbek). 

Tree to 6—10 m high, with broad crown; shoots unarmed, thick, young 
and annotinous shoots pubescent-tomentose; leaves firm, glaucous-green 
on both sides, sparsely appressed-pubescent, more densely so above, 
sometimes subglabrous when adult; lower leaves on fruiting shoots obovate- 
cuneate, often oblong, decurrent along petioles, coarsely incised-dentate 
at the apex or trilobate; upper leaves (3)4.5— 6.5 cm long and broad, the 
blade 3—6 times as long as the petiole, rhomboid or broadly obovate, broadly 
cuneate at base, deeply 5-fid, the middle lobe often trifid or 7-partite; 
lobes usually oblong, usually 3 times as long as broad (2.5—4:1), some 
entire, some with 1—4 large teeth at the apex; specimens with all their 
leaves broadly trilobate have been found exceptionally; inflorescences 
3— 5 cm in diameter, (6)8— 14-flowered, hairy to tomentose; pedicels 3 — 7 cm 
long; sepals broadly triangular, acute or acuminate, recurved in fruit; 
corolla 15— 20 mm in diameter; styles 2 or 3; fruits strongly flattened at the 
poles, 15— 25(28) mm in diameter, yellow, from ferruginous-green to orange- 
yellow, often reddish laterally and with reddish dots; nutlets 3, obtusely 
triquetrous, or else 2, with flat ventral side. Fl. June— July, fr. from 
September. (Plate XXIX, Figure 2). 



5773 332 



At 1,000-1,200